FLORA
MALESIANA
SERIES I - SPERMATOPHYTA
Flowering Plants
Volume 11, part 2
Rosaceae, Amaryllidaceae, Alliaceae
Coriariaceae, Pentastemonaceae, Stemonaceae
LIBRARY
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FLORA MALESIANA
SERIES I — SPERMATOPHYTA
Volume 11 -part 2 -1993
Rosaceae
(C. Kalkman — pp. 227-351)
Amaryllidaceae
(D.J.L. Geerinck — pp. 353-373)
Alliaceae
(J.R.M. Buijsen — pp. 375-384)
Coriariaceae
(B.E.E. Duyfjes — pp. 385-391)
Pentastemonaceae
(B.E.E. Duyfjes — pp. 393-398)
Stemonaceae
(B.E.E. Duyfjes — pp. 399-409)
v. II
CIP-GEGEVENS KONINKLUKE BIBLIOTHEEK, DEN HAAG
Flora
Flora Malesiana. Series I, Spermatophyta : Flowering plants. -
Leiden : Rijksherbarium / Hortus Botanicus, Leiden University
Vol. 11, pL 2: Rosaceae / C. Kalkman. Amaryllidaceae / D.J.L. Geerinck.
Alliaceae / J.R.M. Buijsen. Coriariaceae / B.E.E. Duyfjes. Pentastemonaceae / B.E.E. Duyfjes.
Stemonaceae / B.E.E. Duyfjes. - 111.
Comp. and publ. under the auspices of Foundation Flora Malesiana. - Met index, lit. opg.
ISBN 90-71236-19-6
Trefw.: flora ; Zuidoost-Azie.
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© 1993 Foundation Flora Malesiana
No part of the material protected by this copyright notice may be reproduced or utilized in any form
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Flora Malesiana ser. I, Vol. 11 (2) (1993) 227-351
ROSACEAE
(C. Kalkman, Leiden, The Netherlands)
Rosaceae Juss., Gen. PI. (1789) 196, nom. cons.; Hutch., Gen. Flow. PL 1 (1964)
174-216; Vidal, Fl. Camb., Laos & Vietnam 6 (1968) 1-210 (excl. Rubus); Nguyen
Van Thuan, ibid. 7 (1968) 1-83 (Rubus); Vidal, Fl. Thailand 2 (1970) 31-74 (Rubus by
Nguyen Van Thuan); Tirvengadum, Fl. Ceylon 3 (1981) 328-378. — Type genus:
Rosa L.
Woody or herbaceous plants. Leaves usually spirally arranged, sometimes distichous,
rarely opposite (not in Malesia), simple or compound. Stipules on the twig or on the base
of the petiole, free or adnate to petiole, rarely absent Inflorescences various. Flowers usu-
ally bisexual and actinomorphic. Hypanthium ('calyx tube' of many authors) usually very
distinct, from saucer-shaped to tubular or campanulate, the sepals, petals, and stamens
inserted on its rim, its inside usually lined by a nectariferous disk. Sepals usually 5, free,
in some tribes an epicalyx also present. Petals usually 5, free, from large and showy to
small and not or hardly distinct from sepals, in some genera/ species absent. Stamens
usually numerous, but sometimes the number distinctly related to the number of perianth
leaves, filaments free, anthers bilocular, dehiscing longitudinally. Pistil(s) 1 to many, free
or variously connate with each other and/or with the hypanthium, ovary(ies) superior to
inferior, style(s) present, ovule(s) 1 to several (often 2) per locule, anatropous, ascending
or pendulous. Fruits various, fleshy or dry, dehiscent or not. Seed(s) 1 to several, with-
out or with scanty endosperm, cotyledons fleshy or flat.
Distribution — A large family with worldwide distribution, including more than 3000
species in c. 100 genera. Almost all genera which are represented in Malesia have their ac-
tual centre of distribution in temperate to subtropical regions on the Northern Hemisphere.
Some of those are large or medium large genera with only one or two species in Malesia
(Rosa, Alchemilla, Eriobotrya), others have a more or less distinct sub-centre in the Male-
sian region (Prunus, Rubus, Potentilla). Exceptional is Acaena, a genus with a Southern
Hemisphere distribution, of which one species also occurs in New Guinea.
Kalkman [Bot. J. Linn. Soc. 98 (1988) 37-59] postulated a Southern (Gondwanan)
origin for the family and migration via three routes. Malesia in this view was reached
mainly from the Asian continent (Laurasia), which in turn was reached by way of South,
Central, and North America and via Beringia. Partly maybe the continent was also reached
directly from Gondwana by transport on the Indian 'raft' (Alchemilla?). A third route was
via Australia (Acaena). Most authors, however, favour a Laurasian origin for the family.
Habitat — The majority of Malesian Rosaceae belongs to the mountain flora and occurs
only above 1000 a 1500 m altitude, in montane forest types, thickets or (sub)alpuie grass-
lands. Only in the genus Rubus (a dozen species) and in Prunus (more than 20 species) an
appreciable proportion of the species are (also) found in the lowlands.
(227)
228 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Ecology — As is true for almost all Malesian higher plant families, no autecological
research has been carried out for members of Rosaceae. From the habitats where the spe-
cies have been collected, some superficial conclusions may be drawn about preferences or
tolerances for light, temperature and soil conditions and wherever possible, the paragraphs
on Habitat and Ecology contain this kind of information.
Pollination is undoubtedly normally by (unspecified) insects, as in the European rela-
tives. Apart from the formation of a good quantity of pollen and the secretion of nectar by
the disc, there are no specializations in the flowers related to pollination by specific kinds
of insects. Only for Acaena wind-pollination might be inferred, but experimental or obser-
vational evidence is lacking in literature or on labels.
For dispersal most Rosaceae rely heavily on animals. Exceptions are found in genera
with multi-seeded follicles with dry seeds (in Malesia only Neillia) where dispersal is by
ballistochory. The same is true for most of the Potentilla species that have the dry achenes
in the cups formed by the hypanthium, sepals and epicalyx. Some genera have dry achenes,
imbedded in or surrounded by a fleshy spurious fruit (Rosa and also the not indigenous
Fragaria and Potentilla indica). In these cases the hypanthium, resp. the torus functions as
the attractant for endozoochory by snails, birds, or other animals. Many Rosaceae of dif-
ferent tribes have gone the way to fruits with a fleshy or juicy layer in their walls (drupes,
either single or as collective, or pomes) and obviously these are also endozoochorous.
Epizoochory is only exercised by Acaena and Agrimonia which possess spines on their
hypanthium in which the fruit is included.
Taxonomy and Phylogeny — In modern systems the Neuradaceae and the Chryso-
balanaceae [for the latter, see Flora Malesiana I, 10 (1989) 635-678] are mostly not in-
cluded in Rosaceae, as in earlier classifications, but recognized as families in their own
right.
In the family Rosaceae as implied in the previous paragraph usually four subfamilies
are distinguished: Spiraeoideae, Rosoideae, Maloideae (Pomoideae), and Prunoideae
(Amygdaloideae). The last-mentioned two groups are undoubtedly two end-branches in
the phylogenetic tree, well recognizable, distinct, and natural (holophyletic) taxa. This
cannot be said for the two other subfamilies. The group Spiraeoideae contains the genera
with dry, dehiscent fruits. Dehiscent follicles is a plesiomorphic (primitive) character in
the family and the genera possessing this character should better be included in a taxon
with the genera that have been derived from them. Considering the likeness of the flowers
of some Spiraeoideae and those of some Maloideae like Cotoneaster and Pyracantha, I
would be inclined to enlarge the subfamily Maloideae with at least part of the Spiraeoid
genera.
The Rosoideae are quite heterogeneous and they have probably to be united with other,
'Spiraeoid', genera to form another holophyletic branch, in which maybe some subdivision
is possible.
A phylogenetic analysis, only considering morphological characters [Kalkman, Bot. J.
Linn. Soc. 98 (1988) 37-59] was not successful and should be repeated with an augmented
set of characters, also anatomical and chemical ones. Awaiting this, the recognition of the
four classical subfamilies is hardly justified.
Kalkman — Rosaceae f^l
The next lower level of classification was used by Hutchinson, I.e., who divided the
family in some twenty tribes. Some of these are heterogeneous, some others contain only
one or two 'difficult' genera that have not yet found a good place with their nearest rela-
tives in the phylogenetical sense.
The firm core of a tribal classification consists of the following tribes:
S Spiraeeae (see p. 244)
S Neillieae (see p. 245)
S Gillenieae (maybe to be split into two tribes) (not in Malesia)
R Rubeae (see p. 247)
R Potentilleae (see p. 285)
R Dryadeae (probably to be divided into two tribes) (not in Malesia)
R Poterieae (see p. 297)
R Alchemilleae (usually included in Potentilleae or Poterieae) (see p. 301)
R Roseae (see p. 303)
M Maleae (Pomeae) (see p. 306)
P Pruneae (including Osmaronia?) (see p. 319)
S = tribes belonging to Spiraeoideae in the classical sense; R = Rosoideae; M = Maloi-
deae; P = Prunoideae.
The following genera have not found a natural place in one of the eleven (or thirteen
after dividing two of them) tribes mentioned above:
— the genera composing the tribe Quillajeae in Hutchinson's classification, a rather hetero-
geneous assemblage of Spiraeoid problem cases: Quillaja, Kageneckia, Exochorda,
Lindleya, Vauquelinia, Lyonothamnus;
— a number of lone genera of uncertain disposition as to tribe and often also subfamily:
Holodiscus, Rhodotypos, Kerria, Neviusia, Cercocarpus, Coleogyne, Filipendula,
Potaninia, Adenostoma.
None of the genera mentioned above occur in Malesia.
Morphology — As apparent from the family description, there is variation in many
characters of leaves, flowers, and fruits. The presence of a well-developed hypanthium, a
probably axial outgrowth from the top of the pedicel surrounding the pistil(s), is about the
only character that is common to all Rosaceae. The elaboration of this hypanthium causes
much of the variation in flowers and fruits. The plesiomorphic (original) situation is still
present in Spiraeoid genera that have a small number (up to 5) multi-ovulate ovaries on
the bottom of a cupular hypanthium, the ovaries developing into ventrally dehiscent, dry-
walled follicles containing several seeds.
Adnation of the ovaries to the inside of the hypanthium, accompanied by a more or
less complete fusion of the ovaries with each other, creates the possibility for the evolu-
tion of the fleshy, (semi-)inferior fruits that are typical for Maloideae. In this group the
exocarp of the inferior fruit is certainly hypanthial, the endocarp (membranous to woody)
is certainly carpellary, the more or less fleshy mesocarp may be either or both. In the
descriptions in this treatment the terms exocarp, mesocarp, and endocarp are used in
230 Flora Malesiana ser. I, Vol. 11 (2) (1993)
their topographical sense, for superior as well as inferior fruits and thus not implying a
carpellary origin.
Another line of evolution is the change of dry, multi-seeded follicles into dry, 1 -seeded
achenes, that later may develop a fleshy fruitwall and become drupaceous. Examples are
manifold in Rosoideae (e. g. Rubus with many pistils per flower) and all Prunoideae have
drupes too, with one pistil per flower.
Vegetative Anatomy — Although many papers have been published on the leaf anat-
omy of the Rosaceae, little is known of the anatomy of the tropical representatives, especi-
ally of the Malesian species. The situation for wood anatomy is much better with recent
comprehensive studies by Zhang (1992), Zhang & Baas (1992) and Zhang et al. (1992).
The following is a concise summary for the Malesian representatives (wild and cultivated)
of data surveyed more extensively in Metcalfe & Chalk (1950) and the above-mentioned
wood anatomical studies, amplified with scattered data from the other papers cited below.
Leaf anatomy. Trichomes if present usually unicellular, but tufted or stellate hairs occur
in Potentilla p.p. and Rubus p.p.; stalked capitate glands have been recorded in Alchemilla,
Fragaria, Potentilla, Prunus, Rosa, Rubus, and Sanguisorba. Epidermal cells of lower
epidermis sometimes papillate.
Extrafloral nectaries present on the petiole and various parts of the leaf blade of Prunus
p.p.; leaf teeth glandular or hydathodal in species of Alchemilla, Fragaria, Prunus, Pyrus,
Rubus, Sanguisorba, and Spiraea. Stomata almost always confined to the lower leaf sur-
face, usually anomocytic, but cyclocytic, staurocytic, tetracytic, and actinocytic types may
also occur (Lu et al. 1991). Upper epidermal cells often (partly) mucilaginous. A hypoder-
mis is differentiated in some species of Prunus ('Pygeum') and Rubus (section Micrantho-
batus). Mesophyll dorsiventral. Vascular bundles of minor veins with or without scleren-
chyma in bundle sheath, only rarely vertically transcurrent. Vascular system of midrib and
petiole ranging from a single collateral bundle to more complex, open or closed systems.
Nodes usually trilacunar, but 5-, 7-, and 9-lacunar nodes recorded in Rubus (Kato 1966,
1967). Crystals solitary and/or clustered. Tanniferous cells common. Mucilage idioblasts
present in the mesophyll of some species.
Wood anatomy. Growth rings faint or absent. Vessels diffuse, typically mostly soli-
tary, but radial vessel multiples common in Prunus s.L, and vessel clusters fairly common
in Rubus. Vessel frequency and diameter very variable depending on plant habit and ecol-
ogy. Intervessel pits, nonvestured, alternate, ranging from minute to large (2-12 um);
vessel-ray pits usually similar but half-bordered and slightly smaller (but conspicuously
smaller in Prunus p.p.). Helical wall thickenings present in Rosa, all Old World Maloi-
deae, and Prunus, but usually weakly developed or restricted to vessel element tails in the
Malesian species (as far as studied). Gummy contents common in heartwood vessels.
Ground tissue fibres typically fibre-tracheids with distinctly bordered pits common in
radial and tangential walls, but in Prunus pits mainly confined to the radial walls, and in
some species (belonging to the subgenera Amygdalus and Laurocerasus) also much re-
duced in size so that fibres tend to the libriform type. Parenchyma typically scarce, scanty
paratracheal and apotracheal diffuse; in the Maloideae parenchyma more abundant and
sometimes diffuse-in-aggregates. Irregularly zonate parenchyma bands restricted to Prunus
231
Kalkman — Rosaccae
p p CPygeum 1 and subs. Laurocerasus p.p.). Rays l-3(-4)-seriate in all Maloideae
and in Alchemilla (1-seriate) and Potentilla [l-2(-3)-seriate]; in the remaining genera
(Prunus and most Spiraeoideae and Rosoideae) rays of two more or less distinct sizes,
with the wide rays 3-8(-16)-seriate. Ray composition varying from homocellular in
Micromeles and Photinia p.p.; heterocellular with one to several rows of square to upright
marginal cells in Prunus and most Malesian Maloideae (Kribs' heterogeneous II & III); to
larcely composed of square to upright cells in all shrubby Spiraeoideae and Rosoideae
(so & -called 'juvenilistic rays'). Crystals absent, or present as rhomboidal crystals in ray
cells (Rosoideae and Spiraeoideae) or in, usually enlarged, chambered axial parenchyma
cells (Maloideae); in species of Prunus druses (subg. Amygdalus, Laurocerasus p.p.,
Padus, Prunus, and 'Pygeum') may occur as well as, or instead of rhomboidal crystals
(the latter are found in subg. Laurocerasus and Padus), in ray and/or axial parenchyma
cells. Traumatic gum ducts sometimes present in species of Prunus (except species be-
longing to subg. Prunus).
The wood anatomical diversity of the Rosaceae lends itself well for microscopic wood
identification and a contribution to phylogenetic classification of the family (Zhang' 1992).
The Spiraeoideae and Rosoideae are wood anatomically fairly heterogeneous but insepar-
able; the Maloideae are a very coherent group showing only very limited wood anatomical
variation. Prunus, on the other hand is wood anatomically very diverse and several groups,
largely coinciding with the present subgeneric boundaries, can be distinguished, lending
some support to their treatment as separate genera: Amygdalus, Laurocerasus, Padus,
Prunus s.s., and Pygeum. More detailed leaf and wood anatomical studies of the Male-
sian taxa will certainly yield much information of taxonomic significance.
References- Devadas, C & C.B. Beck, Amer. J. Bot. 59 (1972) 557-567 (nodal anatomy Physocar-
pus Prunus Rubus. Potentilla, Geum). - Kato, N., J. Jap. Bot. 41 (1966) 101-107 & ibid. 42 (1967)
161-168 (nodal anatomy). - Lersten, N.R. & J.D. Curtis, Can. J. Bot. 55 (1977) 128-132 (trichomes
of Rubus, Physocarpus, Fragaria, Potentilla); ibid. 60 (1982) 850-855 (hydathodes and water pores). -
Lu, L.T..Z.L. Wang & G. Li, Cathaya 3 (1991) 93-108 (leaf anatomy of Erio bo try a, Photinia, Rhaphio-
lepis. Sorbus, Stranvaesia). — Metcalfe, C.R. & L. Chalk, Anatomy of the Dicotyledons 1 (1950) 539-
550 _ Morvillez, M.F., Recherches sur l'appareil conducteur foliaire des Rosacees, des Chrysobalana-
cees et de Legumineuses. Thesis, Lille (1919). - Schnell. R., G. Cusset & M. Quenum, Rev. Gen. Bot.
70 (1963) 269-342 (extrafloral nectaries). — Simomura, T. & H. Kurokawa, J. Jap. Bot. 26 (1951)
339-343 (leaf anatomy Prunus). - Singh, V. & D.K. Jahn, Curr. Sci. 44 (1975) 63 (stomatal poly-
morphism in Prunus). - Zhang, S.-Y., Blumea 37 (1992) 81-158 (systematic wood anatomy of the
Rosaceae) — Zhang S.-Y. & P. Baas, IAWA Bull. n.s. 13 (1992) 7-1-91 (wood anatomy of Chinese
Rosaceae). - Zhang, S.-Y., P. Baas & M. Zandee, IAWA Bull. n.s. 13 (1992) 307-349 (ecological
wood anatomy). P - Baas
Palynology — Pollen of representatives of some 80 genera of Rosaceae has been stud-
ied in greater or less detail (see Tissot 1990). A comprehensive family treatment is not
available up to now. Detailed regional accounts are those by Reitsma (1966), Teppner
(1966) and Eide (1981) for NW Europe, and those by Hebdaet al. (1988a, b, 1990, 1991)
for W Canada, and by Naruhashi & Toyoshima (1979) for Japan.
Rosaceae is a stenopalynous family. The pollen grains are isopolar, radially symmetric,
subspheroidal monads. The polar axis (P) is 10-60 urn (mostly 20-35 urn), the equato-
232 Flora Malesiana ser. I, Vol. 1 1 (2) (1993)
rial diameter is 10-50 urn. The Spiraeeae, Gillenieae, Holodiscus, Filipendula and Ade-
nostoma have small pollen (P usually < 20 (irn). The largest grains occur in Agrimonia
and Mespilus (P up to 50-60 urn).
The apertural system is generally tricolporate, though di-, tetra-, syn- and pericolporate
grains may be occasionally found in tricolporate samples. Small grains have often indistinct
or ill-defined endoapertures (colpate, colporoidate). Pollen of Woronowia, a monotypic
genus included in Sieversia by Hutchinson (1964), is 5- (or 6-)colporate (Li 1990). Usu-
ally the colpi are relatively long, but Polylepis and Cliffortia (Poterieae) have brevicolpate
and porate ectoapertures respectively. All Potentilleae have operculate ectoapertures. Oper-
cula occur also in several genera of Poterieae (e.g., Acaena, Agrimonia, Sanguisorba),
and in a number of Rosa species. In Sanguisorba spp. the opercula are as wide as the
mesocolpia, which often led to describing the grains as being 6-colporate. Many rosace-
ous pollen grains show 'pore flaps', i.e. more or less protruding sexinous extensions of
the mesocolpia arching over an endoaperture, and that occasionally may form an equa-
torial bridge (Hebda & Chinappa 1990).
Exine stratification is usually distinct with light microscopy. The scarce electron micro-
graphs published show a columellate infratectal layer. Ornamentation is mostly meridional-
ly striate, but much variation is found in the length, height, width and pattern of the mini and
lumina, and the size and number of the perforations within the lumina. Agrimonia and Sor-
baria pollen is finely transversely striate. Other ornamentation types include rugulate, psilate
(e.g. Cotoneaster), perforate-microreticulate (Neillia) and verrucate-scabrate (e.g., Acaena,
Alchemilla, Sanguisorba). Many intermediate forms exist. The tribe Poterieae is most di-
verse with respect to ornamentation. The scabrate type of Acaena and Polylepis might be
associated with wind-pollination. Pollen of these plants is recorded in many diagrams of
lake sediments from Colombia and Venezuela (Smit 1978; Salgado-Labouriau 1979).
Obviously, Rosaceae pollen offers few possibilities for subdividing the family. The
detailed studies of Canadian Rosaceae pollen allow the recognition of many pollen types
(Hebda et al. I.e.), but future work must reveal whether these types have any systematic
and phylogenetic significance. Very few fossil pollen has been reported, all from the Oli-
gocene onwards (Muller 1981).
The pollen of Chrysobalanaceae and Rosaceae is readily distinguisable, but the differ-
ences are relatively small (Prance 1989). Pollen of the Neuradaceae is clearly different
(see Erdtman 1952; Van Zinderen Bakker & Coetzee 1959).
References: Eide, F., Grana 20 (1981) 101-118. — Erdtman, G., Pollen morphology and plant taxon-
omy (1952). — Hebda, R.J. & C.C. Chinnappa, Rev. Palaeobot. Palynol. (1990) 103-108. — Hebda,
R.J., C.C. Chinnappa & B.M. Smith, Grana 27 (1988a) 95-113; Can. J. Bot. 66 (1988b) 595-612;
ibid. 68 (1990) 1369-1378; ibid. 69 (1991) 2583-2596. — Hutchinson, J., The genera of flowering
plants 1 (1964). — Li, C.-L., Chin. J. Bot. 2 (1990) 150-153. — Muller, J., Bot. Review 47 (1981)
1„142. _ Naruhashi, N. & Y. Toyoshima, J. Phytogeogr. Tax. 27 (1979) 46-50. — Prance, G.T.,
in Flora Malesiana I, 10 (1989) 635-678. — Reitsma, T., Acta Bot. Need. 15 (1966) 290-307. —
Salgado-Labouriau, M.L., Grana 18 (1979) 53-68. — Smit, A., Rev. Palaeobot. Palynol. 25 (1978)
393-398. — Teppner, H., Phyton (1966) 224-238. — Tissot, C, Sixth bibliographic index to the
pollen morphology of Angiosperms (1990). — Zinderen Bakker, E.M. van & J. A. Coetzee, South
African pollen grains and spores 3 (1959) 104-200. n ... ... , TT
^ ^ R.W.J.M. van der Ham
Kalkman — Rosaceae ^
Phytochemistry — General remarks. Chemistry and chemotaxonomy of Rosaceae
were reviewed twice in recent time (Hegnauer 1973, 1990). In these treatises Chrysoba-
lanaceae with their characteristic seed oils were included as a subfamily in Rosaceae. A
series of papers treating the impact of secondary metabolites, predominantly phenolic-
compounds, on the classification and infrafamiliar evolution of Rosaceae was published
by Challice (1973, 1974, 1981), and a short chemotaxonomic discussion of Rosaceae
was included in an essay of Hegnauer (1976). For the present purpose references usually
will only be given for papers not cited in one of the forementioned publications.
Worldwide, many members of Rosaceae are highly esteemed in traditional medicine.
Several recent investigations are concerned with rosaceous medicinal crude drugs, including
some eastern Asiatic ones; they will be mentioned in the present phytochemical summary.
Leaf phenolics are often heavily overrated in chemotaxonomic discussions (Hegnauer
1990: 373-375). They are doubtlessly valuable characters at generic and lower levels,
but they should be used with extreme care only at higher hierarchic levels; tannins and iso-
flavonoids are perhaps the taxonomically most promising phenol classes at suprageneric
levels.
Cyanogenic glycosides. Since more than 160 years amygdalin, the gentiobioside of
mandelonitrile, is known from seeds of bitter almonds, and prunasin, a glucoside of
mandelonitrile, was prepared in 1895 from amygdalin by cleaving off one molecule of
glucose. For a thorough discussion of cyanogenesis and its possible taxonomic meaning
in Rosaceae see Fikenscher et al. (1981). Amygdalin has been detected in seeds of many
species oiPrunus and several genera of Maloideae. In vegetative parts of both taxa amyg-
dalin is usually replaced by prunasin. For a long time these two cyanogenic glycosides,
which release benzaldehyde on hydrolysis, were considered to be characteristic of the
family. This is true, however, only of the genus Rrunus with the basic chromosome num-
ber (x = 8) and of Maloideae (x = 17). True Rosoideae (x = 7) do not produce cyanogenic
compounds. The genera Exochorda, Oemleria and Prinsepia (all with x = 8 and all consid-
ered to be Prunoideae) have weakly cyanogenic leaves and twigs with a still unidentified
cyanogenic glycoside which is not prunasin.
The Spiraeoideae sensu Schulze-Menz 1964 (mostly x = 9) are rather heterogeneous
with regard to cyanogenesis. Prunasin is present in leaves of Gillenia trifoliata, Spiraea
prunifolia (release of HCN and benzaldehyde; not by other Spiraea-iaxa.) and Aruncus
Silvester (also in rootstocks), and Sorbaria-taxa. (possibly also Chamaebatiaria millefolium)
produce heterodendrin and aromatic esters of cardiospermin. Kageneckia with x = 17 pro-
duces prunasin and resembles Maloideae in this respect.
A third type of cyanogenic glycosides, the tyrosine derived dhurrin, was demonstrated
to be present in Cercocarpus and Chamaebatia (both with x = 9), whereas in other 'rosoid'
taxa with x = 9 hitherto unidentified cyanogenic glycosides, which do not release benzal-
dehyde, occur in small amounts: Kerrieae with Coleogyne ramosissima, Kerria japonica,
Neviusia alabamensis and Rhodotypos scandens, and Adenostomeae with Adenostoma
fasciculatum and sparsifolium. Thus Rosaceae use at least three pathways, the phenylala-
nine-route, the tyrosine-route and the leucine-route, for the production of their cyanogenic
glycosides. The leucine-pathway is also known from Mimosaceae, Crassulaceae and Sa-
pindaceae.
234 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Phenolic constituents. All rosaceous plants are accumulators of phenolic and poly-
phenolic constituents, but the profiles of phenolics vary widely with plant parts and with
taxa. Simple phenolic glycosides like arbutin [Pyrus, Sorbaria, Adenostoma, but not Exo-
chorda (Hegnauer 1990: 374)], picein (the glucoside of 4-hydroxyacetophenone; bark of
Amelanchierp.p.), gein (vicianoside of eugenol; Geum s.l. p.p.), sweet-tasting dihydro-
chalcone glucosides phloridzin, sieboldin and trilobatin [Pyrus and according to Challice
(1974) Docynia and Sorbaria, but not Adenostoma] and the glycosidic derivatives of sali-
cylic acid monotropitin and spiraein (Filipendula) are taxonomic markers of the genera
mentioned or at least of a number of their species.
The closely related species Prunus laurocerasus (prunasin) and P. lusitanica (lusitani-
coside, the rutinoside of the monophenolic phenylpropanoid chavicol) can easily be dis-
cerned by their major leaf constituents mentioned, and the glucosidic phloracetophenone
derivative domesticoside was hitherto only isolated from the bark of Prunus domestica.
Mahaleboside is a 5-glucosyloxycoumarin of Prunus mahaleb; coumarin, herniarin and a
number of 5-hydroxylated and O-methylated coumarins, such as tomentin and fraxinol,
seem to be rather characteristic bark constituents of certain Prunus species. Biologically
active acylphloroglucinol derivatives occur in flowers of Hagenia abyssinica, which were
formerly used as taenifugum (kosotoxin, protokosin), and in the Chinese medicinal plant
Agrimonia pilosa (agrimophol and the agrimols A-C).
Lignans (Ayres & Loike 1990) were isolated from Maloideae [9'-xyloside and 9'-rham-
noside of (+)-lyoniresinol: Sorbus aucuparia, Cotoneaster depressus] and from Prunoideae
[prinsepiol, a furofuranoid lignan from Prinsepia utilis and pygeoside, the 9-xyloside of
(-)-lyoniresinol from Pygeum acuminatum]. For aucuparin-like biphenyls see under phyto-
alexins.
Many more genus- or species-characteristic simple phenolic compounds and their gly-
cosides could be listed without difficulties, but at higher taxonomic levels (tribes, subfam-
ilies, family) hydroxybenzoic acids, hydroxycinnamic acids, flavonoids and tannins are
the predominant classes of phenolic constituents.
Bate-Smith (1961, 1962, 1965) showed, that /?-coumaric andcaffeic acid (= 4-hydroxy-
and 3,4-dihydroxycinnamic acid), the flavonols kaempferol and quercetin and cyani-
din generated from procyanidins (condensed tannins) occur widely in rosaceous leaf hy-
drolysates, that trihydroxylation of the B-ring of flavanoid compounds (e.g. myricetin,
prodelphinidins) is rare, and that ellagic acid indicating presence of ellagitannins is re-
stricted to true Rosoideae (x = 7). Moreover, Bate-Smith already noted incidental pres-
ence of flavones (apigenin, luteolin) and 6-hydroxylated flavonoids (quercetagetin) in the
family.
The screening for rosaceous leaf phenolics was much extended by Challice. He showed
general occurrence of 3-caffeoylquinic acid (chlorogenic acid) and restriction of mixtures
of dicaffeoylquinic acids, known as isochlorogenic acid, to many genera of Maloideae,
inclusive of sections Aria and Aucuparia of Sorbus and of Lindleya. At this point the non-
phenolic cinnamic acid should be mentioned. It occurs in large amounts in hydrolysates of
several species of Spiraea; originally it is present as cinnamoyl-p-glucopyranose and an
acylated derivative, spirarin. Benzoylglucose was isolated from Luetkea pectinata. Challice
73S
Kalkman — Ros aceae til
stressed the taxonomic importance of flavone-C-glycosides within Maloideae, including
Dichotomanthes, and the rather sporadic occurrence of these metabolites elsewhere in the
family.
As already mentioned, the presence of isoflavones in Rosaceae may be taxonomically
rewarding, because they possibly indicate affinities with Leguminosae [see for isoflavones
the treatment of Mlmosaceae in Flora Malesiana 11 (1), p. 19]. The isoflavones genistein,
prunetin, biochanin-A and their glucosides prunitrin (prunetin-4'-glucoside), prunetino-
side (prunetin-5-glucoside) and biochanin-A-7-glucoside have not yet been traced as leaf
constituents; they were isolated from wood and bark of several species of Primus, from fruit
stalks of Prunus avium and P. cerasus and from flowers and fruits of Cotoneaster pannosa
and serotina. Many more peculiar flavonoids including flavanones (e.g. Bilia et al. 1991),
flavanonols (e.g. Yoshida et al. 1989a), the 8-methoxyflavonols sexangularetin and cor-
niculatusin which occur in Dryas octopetala, Cowania mexicana and Purshia glandulosa
(all Dryadeae with x = 9), many more O-methylated flavonoids, tricetin, a flavone with
a 3',4',5-trihydroxylated B-ring (Luetkea pectinata), and even 2-phenoxychromones
(Hashidoko et al. 1991a) are produced by Rosaceae. For more information about the
multiformity of flavonoid metabolism in the family and its possible taxonomic meaning
see Challice (1981) and Hegnauer (1973, 1990). The most conspicuous rosaceous
hydroxybenzoic acid is gallic acid (3,4,5-trihydroxybenzoic acid); it will be mentioned
under tannins.
Tannins (compare also Mimosaceae treatment). Proanthocyanidins (condensed tannins)
seem to be more of less ubiquitous in leaves, flowers, fruits, stems and roots of Rosa-
ceae; they are accompanied by their monomelic building stones (+)-catechin and (-)-epi-
catechin and many (4-8)- and (4-6)-linked catechin dimers and trimers with low tanning
activity; the proanthocyanidins with strong tanning action are assumed to be usually tetra-
mers and higher oligomers.
Double-linked A-type procyanidins are known from the bark of Rhaphiolepis umbel-
lata and from Prunus spinosa. (+)-Gallocatechin and prodelphinidins which are rarely
present in leaves seem to occur more often in 'tannin '-fractions of fruits (Maloideae p.p.)
and roots (Sanguisorba officinalis, Potentilla erecta). Purshia tridentata and Coleogyne
ramosissima vielded about 39c of true condensed tannins (average mol.wt. 13-1400. i.e.
tetra- to pentamers) from winter dormant twigs of current season growth; the tannins of
the two taxa differed in stereochemistry and biological activity (Clausen et al. 1990). The
Purshia tannin was found to have a catechin/epicatechin-ratio of about 55 : 45 and to be
prefered by snowshoe hares in a choice feeding bioassay to the Coleogyne tannin, which
is predominantly based on epicatechin. This shows that the always highly complex con-
densed tannin-fractions may have an extremely diverse spectrum of biological activities
which depend on hydroxylation patterns and stereochemistry of their building stones, on
the nature and stereochemistry of interlinkages in the oligo- and polymers and on degrees
of polymerisation.
In true Rosoideae (x = 7) the situation is even more complex, because in these plants
condensed tannins are accompanied by gallo- and ellagitannins. and because catechins may
also be linked with other aromatic metabolites; the pilosanols-A to -C are antimicrobial
236 Hora Malesiana ser. I, Vol. 11 (2) (1993)
compounds of Agrimonia pilosa in which C-8 of (-)-epicatechin is combined via a methy-
lene group with acylphloroglucide residues (Kasai et al. 1992). In recent times hydroly-
sable tannins were thoroughly investigated for a number of medicinally used crude drugs.
Examples from Rosaceae are roots of Rosa davurica (Yoshida et al. 1989a, 1991), hips
and fresh leaves of Rosa laevigata (Yoshida et al. 1989b), petals of Rosa rugosa (Hatano
et al. 1990), petals of 'apothecary's rose' (Eugster & Marki 1991), Alchemillae Folium
(mainly Alchemilla xanthochlora), which seems to contain only hydrolysable tannins
(Geiger 1991), and Tormentillae Radix which is rich in condensed tannins, but also con-
tains ellagitannins (Geiger 1991) and which derives from Potentilla erecta.
Root cultures of Sanguisorba officinalis yielded gallic acid, (+)-catechin, (+)-galloca-
techin, procyanidin-B3, three gallotannins (2,8%) and the ellagitannins pedunculagin,
sanguiin-H6 (up to 5,9%) and sanguiin-Hll (up to 2,3%) and 4,6-hexahydroxydiphe-
noylglucose (Ishimaru et al. 1990). Strawberries (Fragaria x ananassa cv. Kent) and
raspberries (Rubus idaeus) contain small amounts of ellagitannins, and casuarictin was
isolated from strawberries (Daniel et al. 1991).
Lamaison et al. (1990) investigated 42 rosaceous taxa representing all four traditional
subfamilies for tannin content and observed a range from 1,7 (flowers of Kerria japonica)
to 25,1% (roots of Potentilla erecta); they demonstrated that biological activity measured
by inhibition of the pancreatic endopeptidase enzyme elastase is not correlated with tannin
concentration. Most active tannins were found in flowers and leaves of Alchemilla xantho-
chlora, flowers of Filipendula ulmaria, aerial parts of Gewn montanum and G. rivale and
leaves of Sanguisorba minor, all true Rosoideae with x = 7 and usually containing both
condensed tannins and hydrolysable gallo- and ellagitannins. This is another indication for
the pluriformity of biological activities of individual tannin components. Finally it should
be mentioned that trimethylellagic acid was isolated from rhizomes of Sanguisorba offici-
nalis, because ellagic acid methylethers are usually conceived as taxonomic markers of
Myrtales; they also occur in Euphorbiaceae.
A most recent publication of Okuda et al. (1992) confirms confinement of ellagitannins
to Rosoideae with x = 7, including Filipendula (!); moreover, oligomeric ellagitannins are
assumed to show a genus- specific distribution: sanguiin-H6 and -Hll in Sanguisorba and
Rubus, gemin-A in Geum s.l., agrimoniin in Agrimonia, Fragaria and Potentilla and ru-
gosin-D in Filipendula; this publication considers leaf tannins only. For fruits, restriction
of ellagitannins to Rosoideae was shown by Foo & Porter (1981).
Sugars and hexitols. Saccharose is present in all Rosaceae in appreciable amounts; it is
an easily metabolised temporary carbohydrate reserve and is used for the transport of car-
bohydrates. In Prunoideae, Maloideae, most Spiraeoideae in traditional circumscription and
in Kerrieae and Adenostomeae part of saccharose is replaced by the hexitol sorbitol (= gluci-
tol) which got its name from Sorbus aucuparia, one of the first and best sources of this sugar
alcohol. Lack of appreciable amounts of sorbitol is a character of the ellagitannin-produc-
ing true rosoids inclusive of Dry adeae-Geinae, all with x = 7. In this respect part of Dry a-
deae sensu Schulze-Menz (1964), i.e. Dryadinae and Cercocarpinae both with x = 9, seem
to agree better with true Rosoideae (x = 7) than with Spiraeoideae and Kerrieae with x = 9.
Waxes and other lipids. Rosaceous cuticular waxes of leaves and fruits (Maloideae)
usually are rich in free pentacyclic triterpenic acids. Ursolic and oleanolic acid and a num-
Kalkman — Rosaceae ^]_
ber of hydroxylated derivatives (e.g. maslinic and pomolic acid) were isolated from
leaves and pomes of many taxa. Barks of Rosaceae often contain free and/or esterified
pentacyclic triterpenic alcohols and ketones; lupeol, betulin, 23-hydroxybetulin (sorbicor-
tol-B), taraxerol, alnusenol (= glutinol), friedelanol, alnusenone (= glutinone), friedelin
and others were isolated from several taxa. Moreover, barks also contain free and esteri-
fied triterpenic acids; examples are pyracrenic acid (betulinic acid-3-coumarate) from Pyra-
cantha crenulata and betulinic and 3-epibetulinic acid from Spiraea cantoniensis. Most non-
glycosylated bark triterpenes are probably constituents of cork waxes. Besides triterpenes,
lipid fractions of all plant parts contain alkanes, alkenes, alkanols, long-chain fatty acids
and mixtures of phytosterins.
Saponins and pseudosaponins. Saponins are widespread in the family. The sapogenins
are usually derivatives of pentacyclic triterpenes, mostly ursolic acid, but sometimes olea-
nolic or betulinic acids. Three types of sugar attachment to the sapogenins occur: 3-glyco-
sides, esters of the 28-carboxyl group, and the bisdesmosidic saponins which have both
linkages. Compounds which have only the ester-linkage were called pseudosaponins by
French authors, because their properties are different from those of 3-glycosides (true
saponins); tormentol or tormentoside (= rosamultin) is such a pseudosaponin which oc-
curs in many true Rosoideae (x = 7) and in some Maloideae, but could not be detected
neither in Prunoideae and Spiraeoideae nor in Kerrieae; it is the 28-COOH glucose ester
of tormentillic (= tormentic) acid (2a,19a-dihydroxyursolic acid) and seems to combine
easily with condensed tannins to antibiotically active adducts.
As a rule saponins are complex mixtures of closely related compounds. Since a long
time quillajasaponin is known; it occurs in the bark of Quillaja saponaria which is com-
mercially available (Soap bark) and is (was) mainly used as a non-ionic detergent (soap
substitute). Presently quillajasaponin is known to be a mixture of bisdesmosidic com-
pounds of complex structure which have quillaic acid (= 16-hydroxy-23-oxo-oleanolic
acid) as sapogenin and are acylated in the sugar part by two molecules of 3,5-dihydroxy-
6-methyloctanoic acid (a normonoterpenoid compound).
The Japanese crude drug Sanguisorbae Radix (= 'Ziyu') gathered from Sanguisorba
officinalis yielded the 3-arabinoside of pomolic acid (ziyu-glycoside-II) and its bisdes-
mosidic derivative ziyu-glycoside-I which has its 28-carboxyl esterified with glucose.
In recent time the pseudosaponins of leaves of Japanese Rubus-taxa. were investigated
thoroughly. The 28-COOH glucose esters of the ursolic acid derivatives acuminatic
(= euscaphic) acid, 19-hydroxyasiatic (= 23-hydroxy tormentillic) acid and the 3-epimer
of 19-hydroxyasiatic acid could be isolated from R. microphyllus ('Niga-ichigo'; yielded
the niga-ichigosides-Fl to -F3), R. trifidus ('Kaji-ichigo'; yielded the kaji-ichigosides-Fl
and -F2), R. koehneanus (niga-ichigoside-Fl and -F2) and R. x medius, a trifidus hy-
bride (niga-ichigoside-F2 and kaji-ichigoside-Fl). Moreover, roots of the Chinese R. sua-
vissimus contain niga-ichigoside-Fl and suavissimoside-Rl, which is a 28-COOH glu-
cose ester of a derivative of 19-hydroxyasiatic acid (23-CH20H oxidized to 23-COOH).
Niga-ichigoside-Fl was also isolated from Geiun japonicum and from hips of Rosa
sterilis. Russian Sanguisorba minor s.l., i.e. Poterium lasiocarpum and P. polygamum,
yielded a pseudosaponin with caccigenin (2a,21p\23-trihydroxyoleanolic acid) as sapo-
senin.
238 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Recent investigations of leaves of Eriobotrya japonica (Liang et al. 1990; De Tommasi
et al. 1992), Potentillafruticosa (Ganenko & Semenov 1989), hips of Rosa davurica
(Kuang et al. 1989), fruits of Rubus coreanus, crataegifolius and parvifolius (Ohtani et al.
1990), whole plants of Rubus ellipticus (Pal et al. 1991) and root bark or roots and aerial
parts of Sarcopoterium spinosum, Sanguisorba minor s.l. and officinalis, and species of
the endemic Canary Island genera Bencomia, Marcetella and Dendriopoterium (Reher
et al. 1991; Reher 1991) for pentacyclic triterpenes and their pseudosaponins and saponins
yielded taxonomically remarkable results. For instance, Sanguisorba officinalis which
only contains the ziyu-glycosides-I and -H and some recently detected other derivatives of
pomolic acid (Cheng & Cao 1992) is distinct from all investigated members of subgenus
Poterium, Sarcopoterium spinosum and the Canary Island endemics, which all produce
23-hydroxytormentillic acid and its 28-ester glucoside (= niga-ichigoside-Fl).
Rubus coreanus is distinct from other eastern Asiatic species of Rubus by having 0.14-
0.25% coreanoside-Fl, a dimeric pseudosaponin, in leaves and fruits (Ohtani et al. 1990).
Leaves and fruits of Rubus foliolosus do not contain pseudosaponins, but a mixture of
goshonosides (see sub diterpenes; Ohtani et al. 1990). The triterpene (and sesquiterpene)
glycoside profiles of leaves of Eriobotrya japonica growing in China are different from
those of plants growing in Italy (Liang et al. 1990; De Tommasi et al. 1990, 1991, 1992).
Rosaceae also produce tetracyclic dammarane- and cucurbitane-type triterpenes. They
were detected in leaves and twigs of Cowania mexicana (dammarenediol-II) and Cercocar-
pus intricatus (isofouquierol) and in fruits of Purshia tridentata (bitterbrush; the intensely
bitter cucurbitacins-D and -I; occur possibly also in other parts of the plant); all three taxa
belong to Dryadeae-Cercocarpinae and -Purshiinae with x = 9.
Diterpenes. Isoprenoid C20-compounds seem to be rather rare in the family. They oc-
cur in Spiraea japonica and koreana as atisane-type diterpene alkaloids (spirasines, spira-
mines, and others). In the genus Rubus several eastern Asiatic species were shown to
produce large amounts of C20"glycosides in leaves instead of the usual pseudosaponins.
The Chinese Rubus suavissimus contains the intensely sweet-tasting kauranoid steviol-
13,18-bisglucoside rubusoside. From R. chingii, a species also occurring in Japan, where
it is called 'Gosho-igicho', non-sweet labdanoid mono- and bisglucosides, the goshono-
sides-Fl to -F5, were isolated; such diterpenes also occur in fruits of the respective spe-
cies (Ohtani et al. 1990), whereas their roots contain pseudosaponins (e.g. R. suavis-
simus).
Seed reserves. Rosaceae store mainly proteins and fatty oils in their seeds; starch is
absent. The seed oils belong to a 'normal' type with oleic and linoleic acids as main fatty
acids; saturated acids (mostly palmitic) usually approximate 10-15%. Some species of
temperate regions (Filipendula ulmaria, Sanguisorba minor, Rosa p.p.) have linolenic acid
as a third main fatty acid. Reher (1991) found a 18:3/18:2-ratio of 1.4-2.4 in Poten-
tilleae and pf 0.4-1.0 in Sanguisorbeae. Some species of Prunus, notably P. africana
(= Pygeum africanum), P. mahaleb, spinulosa, undulata and yedoensis and others, devi-
ate from the patterns mentioned by having octadeca-9,1 1,13-trienoic (= elaeostearic) acid
as a main fatty acid and resemble in this respect Chrysobalanaceae.
Miscellaneous. Rosaceae produce and store many more classes of metabolites. Exam-
ples are:
Kalkman — Rosaceae ^
(a) Non-volatile organic acids, such as the ubiquitous malic, citric and succinic acids,
and ascorbic acid (= vitamin C) which is present in large amounts in the hips of many
species of Rosa. Isocitric acid which is seldom present in appreciable amounts in plants is
stored in leaves of most investigated species of Rubus.
(b) Some species with glandular hairs produce essential oils containing mosdy mono-
and sesquiterpenoid constituents. Such essential oils are also deposited in the wood of
certain species of Prunus. The best known 'volatile oils' of Rosaceae are rather products
of hydrolysis of glycosides than true essential oils and usually consist for over 90% of
one or two compounds, e.g. bitter almond oil (benzaldehyde from prunasin and amyg-
dalin), methylsalicylate and salicylic aldehyde (from monotropitin and spiraein), eugenol
(from gein), chavicol (from lusitanicoside) and Sorbus aucuparia fruit oil, which consists
of antibiotically active parasorbic acid, the lactone of 5-hydroxy-2-hexenoic acid (= 2-
hexene-5-olide); parasorbic acid is not present as such in the bitter fruits and seeds of
Sorbus species of section Aucuparia, but as the glucosidic bitter precursor parasorboside,
which is 3-glucopyranosyloxy-5-hexanolide, and seems to be a chemical marker of Sor-
bus section Aucuparia. The very expensive true oil of rose is produced from fragrant
flowers of several taxa of Rosa and contains predominantly the monoterpenic alcohols
citronellol, geraniol and nerol and appreciable amounts of phenylethylalcohol; in fresh
young petals these alcohols are present as glycosides. Glycosides of alcoholic mono- and
sesquiterpenes seem to be rather common in the family; some recent examples are leaves
of Eriobotrya japonica (De Tommasi et al. 1990, 1992) and leaves of Spiraea cantoniensis
(Takedaetal. 1990).
(c) Characteristic constituents of fruit aromas, such as raspberry (Rubus idaeus), straw-
berry (Fragaria), quince (Cydonia oblonga), apples (Malus) and cherries (Prunus); in
fresh fruits glycosidic precursors may be present.'
(d) Nitrogen-containing constituents like the proline derivatives of Malus and other
Maloideae and the amines present in the foetid flower smell of some rosaceous taxa (e.g.
Crataegus p.p., Sorbus p.p.). According to Strack (1990) Rosaceae are characterized by
the production of N,N,N-tricoumaroylspermidine in flowers, especially in their androecia.
From a taxonomic point of view metabolites mentioned sub (a) to (d) are unimportant
at suprageneric levels, if the triacyl derivatives of spermidine are excluded. It seems there-
fore to be more rewarding to finish this short chemical survey with a few remarks on phy-
toalexins and recent publications on Prunus constituents.
Phytoalexins. Phytoalexins are antibiotically active compounds produced by plants
after stimulation by infections or similar stresses. Phytoalexins became known from Ro-
saceae only recently. The chemical nature of phytoalexins produced by a taxon depends to
some extent on the triggering agents and the plant parts. Nevertheless some taxonomically
interesting trends can be discerned in Rosaceae. Maloideae tend to produce aromatic phy-
toalexins based on the biphenyl and benzofuran skeleton. The biphenyls aucuparin, 4'-
methoxyaucuparin and rhaphiolepin are produced in infected sapwood or bark of Malus
pumila, Eriobotrya japonica and in stressed leaves of Eriobotrya japonica and Rhaphio-
lepis umbellata, and the biogenetically related benzofurans a-,p- and v-pyrufurans, coto-
nefuran and eriobofuran were extracted from infected sapwood of Pyrus communis and
240 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Cotoneaster lacteus and from diseased leaves of Eriobotrya japonica (Kemp & Burden
1986; Watanabe et al. 1982, 1990; Miyakado et al. 1985). Phenylpropanoid sapwood
phytoalexins are the coumarin scopoletin of Prunus domestica and the lignan iso-olivil
from Prunus jamasakura (Kemp & Burden 1986). Benzoic acid was shown to be the anti-
fungal compound produced after infection by Nectria galligena in apples of cv. Bramley's
Seedling; it can prevent or retard fruit rotting during storage (Swinburne 1973).
Usually the production of phytoalexins is connected with necrosis of attacked cells.
Sapwood phytoalexins are comparable to compounds present in heartwoods which only
contain dead wood parenchyma cells. Aucuparin, for instance, occurs in heartwood of all
investigated species of Sorbus sect. Aucuparia. Moreover, what is known as a phytoalexin
from one plant part may be a normal constituent of perfectly healthy tissues of another part
of the same plant or of other plants, e.g. the coumarin scopoletin, the lignan iso-olivil
and benzoic acid. Lastly, the definition of phytoalexin is rather vague; small amounts of a
given phytoalexin of a given taxon may be present in its healthy tissues. Therefore trigger-
ing of intensified synthesis of a compound by stress is included by some authors in the
phytoalexin concept. Scopoletin in bark of Prunus domestica, coumarin and biogenetically
related compounds in leaves of Prunus mahaleb and glycosides of gentisic acid in wood
and bark of Prunus yedoensis are examples of 'phytoalexins' which are already present in
small amounts in healthy plant parts. Chondrostereum purpureum infection induced not
only synthesis of aucuparin, but also of 2-dehydrotormentillic acid in sapwood of Malus
pumila (Kemp et al. 1985).
Isoprenoid phytoalexin-like compounds were also isolated from damaged leaves of
Rosa rugosa (Hashidoko et al. 1989); they were shown to be watersoluble sesquiterpenes
with the carotane skeleton and named rugosal-A (strongly fungitoxic) and rugosic acid-A
(scarcely fungitoxic); both are monohydroxy-endoperoxides which bear an aldehyde resp.
carboxyl group. Later the same authors reported, that leaf tissues contain a labile precur-
sor, carota-l,4-dienealdehyde which on autoxidation yields rugosal-A and rugosic acid-A
(1990), and that leaves additionally produce many more carotane type sesquiterpenes to-
gether with acaranoid and bisabolanoid oxigenated Cis-compounds (1991b, c). Finally
Hashidoko et al. (1992) observed that rugosal-A and rugosic acid-A are present in the
exudate of the glandular trichomes of Rosa rugosa leaves. The last mentioned observa-
tions suggest that rugosal-A is not a true phytoalexin, but a compound generated by aut-
oxidation from genuinely present precursors.
Recent phytochemical investigations with Prunus-taxa:
Subg. Prunus — Prunus spinosa: Phenolics of flowers (Kolodziej et al. 1991), fruits
(Ramos & Macheix 1990) and branches (Crespo Ibizar et al. 1992; Gonzalez et al. 1992).
Subg. Amygdalus — Prunus davidiana: (+)-Catechin and two flavanone glycosides,
prunin and hesperetin-5-glucoside, from stems (Choi et al. 1991).
Subg. Cerasus — Prunus avium and cerasus: Comparative investigations of inner bark
and seedlings for flavonoids, and detection of prunetin-5-glucoside and tectochrysin-5-
glucoside as chemical markers of P. cerasus and of dihydrowogonin-7-glucoside and
chrysin-7-glucoside as main flavonoids of P. avium: both species have genistein-5-gluco-
side (Geibel et al. 1990, 1991). Prunus serrulata Lindl., a cultigen, yielded 6-caffeoyl-
glucopyranoside and 1,6-dicaffeoylglucopyranoside (Ali et al. 1989). Its var. spontanea
Kalkman — Rosaceae 241
(= P . jamasakura, the wild Japanese mountain cherry) has bitter fruits with prunasin as
main bitter principle (Shimazaki et al. 1991); catechins, sakuranetin-5-glycosides, and the
lignanoid compounds sakuraresinol, dihydrobuddlenol-B and racemic lyoniresinol were
isolated from its bark (Yoshinari et al. 1990). Fruits of P. maximowiczii are also bitter,
they contain bitter terra- to hexaacylsucroses (acyl = one paracoumaroyl + three to five
acetyl residues), epicatechin and a little mandelic acid, but no prunasin (Shimazaki et al.
1991).
Subg. Padus — Fresh bark of P. buergeri yielded no flavonoids, but mono- and bi-
acylated glucopyranoses (caffeic and paracoumaric acid), the 6-caffeoylglucoside of me-
valonolactone, and a little grayanin (Shimomura et al. 1988, 1989a). Bark of P. grayana
is also free of flavonoids, but contains several caffeic, coumaric and 3,4,5-trimethoxy-
benzoic acid esters of glucopyranose, the grayanosides-A and -B and the strongly bitter
grayanin which is prunasin with OH-6 of its glucose acylated by caffeic acid (Shimomura
et al. 1987). Heartwood of P. grayana yielded taxifolin, dehydrodicatechin-A, the salicin
derivatives populin, henryoside and pruyanaside-B and the complex salicylic acid deriva-
tives virgaureoside and pruyanaside-A (Shimomura et al. 1989b). Leaf wax of P. grayana
contains the antioxidative prunusols (Osawa et al. 1991). The bitter barks of P. padus and
P. ssiori contain catechins, bitter prunasin and bitter lignanxylosides lyoniside and ssiori-
side, and schisandriside in P. ssiori and prupaside in P. padus; moreover, P. ssiori yielded
syringin and glucosyringic acid, and P. padus melilotoside and bitter tetra- and penta-
acylated sucroses (Yoshinari et al. 1989, 1990).
Subg. Laurocerasus — From green fruits off. laurocerasus Weinges et al. (1991) iso-
lated the primveroside of benzylalcohol.
The Prunw^-investigations just mentioned demonstrate clearly the enormous intrage-
neric variation of phenolic profiles and suggest once again that most phenolics are taxo-
nomically useful above all at species, section and genus levels.
Summarizing it can be stated that secondary metabolites and seed oils can help a lot to
arrive at a satisfying infrafamiliar classification, but procure only vague indications con-
cerning family affinities [Chrysobalanaceae, Crassulaceae (phenolics, leucine-derived cya-
nogenic compounds), Leguminosae, Sapindaceae].
References and Remarks: Ali, A. A., et al., Pharmazie 44 (1989) 734-735. — Ayres, D.C. & J.D.
Loike, Lignans: chemical, biological and clinical properties, Cambridge Univ. Press (1990). — Bate-
Smith, E.C., J. Linn. Soc., Bot. 58 (no 370) (1961) 39-54 (Potentilla, Prunus); ibid. 58 (no 371)
(1962) 95-173 (Rosaceae: 129-131); Phytochemistry 4 (1965) 535-599 (Quillajeae). — Bilia, A.R., et
al., Phytochemistry 30 (1991) 3830-3831 (hexaacetylpyracanthoside). — Challice, J.S., Phytochemistry
12 (1973) 1095-1101 {Pomoideae); Bot. J. Linn. Soc. 69 (1974) 239-259 (Pomoideae); Preslia (Praha)
53 (1981) 283-304 (Maloideae). — Cheng, Dong-Liang & Cao, Xiao-Ping, Phytochemistry 31 (1992)
1317-1320. — Choi, J.S., et al., J. Nat. Prod. 54(1991)218-224; Planta Medica 57 (1991) 208-211.
— Clausen, T.P., et al., J. Chem. Ecol. 16 (1990) 2381-2392. — Crespo Ibizar, A., et al., J. Nat, Prod.
55 (1992) 450-454. — Daniel, E.M., et al., J. Nat. Prod. 54 (1991) 946-952 (release of ellagic acid
from ellagitannins in vegetable food may be a factor preventing carcinogenesis: this is another example
showing the diversity of biological activities of vegetable tannins, compare Clausen et al. (1990); Lamai-
son etal. (1990)). — De Tommasi, N.. et al., J. Nat. Prod. 53 (1990) 810-815 (sesquiterpene glycosides):
see also Planta Medica 57 (1991 ) 414-416 (hypoglycemic effect of leaf sesquiterpenes and triterpenes>:
J. Nat. Prod. 55 (1992) 1025-1032, 1067-1073 (new sesqui terpen ic glycosides and esterified tnterpenes). —
242 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Eugster, C.H. & E. Marki-Fischer, Angew. Chem. Int. Ed. Engl. 30 (1991) 654-672 (chemistry of rose
pigments). — Fikenscher, L.H., et al., Planta Medica 41 (1981) 313-327. — Foo, L.Y. & L.J. Porter,
J. Sci. Food Agric. 32 (1981) 711-716 (structure of tannins). — Ganenko, T.V. & A. A. Semenov, Khim.
Prirod. Soedin. (1989) 856 (isolation of acids). — Geibel, M., et al., Phytochemistry 29 (1990) 1351-
1353; 30 (1991) 1519-1521. — Geiger, C., Ellagitannine aus Alchemilla xanthochlora Rothmaler und
Potentilla erecta (L.) Raeuschel, Thesis Albert Ludwigs-Univ. Freiburg im Breisgau (1991) (see also
C. Geiger & H. Rimpler, Planta Medica 56 (1990) 585). — Gonzalez, A.G., et al., Phytochemistry 31
(1992) 1432-1434 (A- type procyanidins). — Hashidoko, Y., et al., Phytochemistry 28 (1989) 425-430
(antimicrobial sesquiterpene from Rosa rugosa leaves); ibid. 29 (1990) 867-872; ibid. 30 (1991a) 3837-
3838 (flavone from leaves of Rosa rugosa); Z. fiir Naturforsch. 46c (1991b) 349-356, 357-363 (six bis-
aborosaols from Rosa rugosa); Phytochemistry 30 (1991c) 3729-3739; Agric. Biol. Chem. 55 (1991c)
1049-1053 (rugosic acids); Phytochemistry 31 (1992) 779-782 (rugosal and related sesquiterpenes). —
Hatano, T., et al., Chem. Pharm. Bull. 38 (1990) 3308-3313, 3341-3346 (trigalloylglucoses and ellagi-
tannins). — Hegnauer, R., Chemotaxonomie der Pflanzen 6 (1973) 84-130, 727-730, 784; ibid. 9 (1990)
369-405; Nova Acta Leopoldina, Suppl. no 7 (1976) 45-76; Rosaceae: 57-62 (accumulation of second-
ary products and its significance for biological systematics). — Ishimaru, K., et al., Phytochemistry 29
(1990) 3827-3830. — Kasai, S., et al., Phytochemistry 31 (1992) 787-789. — Kemp, M.S. & R.S.
Burden, Phytochemistry 25 (1986) 1261-1269 (phytoalexins and stress metabolites in sapwood; refer-
ences). — Kemp, M.S., et al., J. Chem. Res. (S) (1985) 154-155. — Kolodziej, H., et al., Phyto-
chemistry 30 (1991) 2041-2047 (5 A-type proanthocyanidins). — Kuang, Hai-Xue, et al., Chem. Pharm.
Bull. 37 (1989) 2232-2233 (Chinese crude drug 'Cimeiguo' yielded 9 triterpenic acids). — Lamaison,
J.L., et al., Ann. Pharm. Franc. 48 (1990) 335-340 (tannin contents and inhibitory activity). — Liang,
Zhou-Zhong, etal., Planta Medica 56 (1990) 330-332. — Miyakado, M., et al., J. Pesticide Sci. 10 (1985)
101-106 (eriobofuran). — Ohtani, K, et al., Phytochemistry 29 (1990) 3275-3280 (dried fruits = Korean
crude drug 'Bog-bun-ja'). — Okuda, T., et al., Phytochemistry 31 (1992) 3091-30%. — Osawa, T., et al.,
Agric. Biol. Chem. 55 (1991) 1727-1731 (prunusols). — Pal, R., et al., Indian J. Chem. 30B (1991)
292-293 (saponins). — Ramos, T. & J.J. Macheix, Plantes Med. Phytotherapie 24 (1990) 14-20
(anthocyanins, caffeoylquinic acids and quercetin glycosides). — Reher, G., Planta Medica 57 (1991),
Sonderheft A76-77 (triterpenoid and fatty acid pattern of several genera oiRosoideae). — Reher, G., et al.,
Planta Medica 57 (1991) 506 (tri terpenoids). — Schulze-Menz, G.K., Rosaceae, in A. Engler, Syllabus
des Pflanzenfamilien 2, 13th ed. by H. Melchior (1964) 209-218. — Shimazaki, N., et al., Phyto-
chemistry 30 (1991) 1475-1480. — Shimomura, H., et al., Phytochemistry 26 (1987) 249-251, 2363-
2366; ibid. 27 (1988) 641-644; Chem. Pharm. Bull. 37 (1989a) 829-830; Phytochemistry 28 (1989b)
1499-1502. — Strack, D., Phytochemistry 29 (1990) 2893-2896. — Swinburne, T.R., in R.J.W. Byrde
& C.V. Cutting (eds.), Fungal pathogenecity and the plant's response (1973) 365-382. — Takeda, Y., et al.,
Phytochemistry 29 (1990) 1591-1593 (monoterpene glucosides). — Watanabe, K, et al., Agric. Biol.
Chem. 46 (1982) 567-568 (aucuparin); ibid. 54 (1990) 1861-1862 (rhaphiolepin). — Weinges, K., etal.,
Liebig's Ann. Chem. (1991) 703-705. — Yoshida, T., et al., Phytochemistry 28 (1989a) 2177-2181;
Chem. Pharm. Bull. 37 (1989b) 920-924; Phytochemistry 28 (1989b) 2451-2454; ibid. 30 (1991) 2747-
2752. — Yoshinari, K, et al., Chem. Pharm. Bull. 37 (1989) 3301-3303 (lignan xylosides); ibid. 38
(1990) 415-417 (Prunus padus); Phytochemistry 29 (1990) 1675-1678 (Prunus jamasakura).
R. Hegnauer
Uses — The usefulness of the Rosaceae is mainly to be found in the presence of many
edible, and often delicious fruits. See: Eriobotrya, Fragaria, Malus, Prunus, Rubus, Pyrus.
Timber hardly enters the world market, but may well be useful on the local scale.
Medicinal uses appear to be scarce in the region judged from label data and Malesian
literature. These sources may not give a complete picture considering the extensive use
made of Rosaceous species in traditional medicine in East Asia (see also the chapter on
phytochemistry, p. 233).
Kalkman — Rosaceae 243
KEY TO THE GENERA
la. Leaves simple, entire or lobed 2
b. Leaves compound, pinnate, palmate, or 3-foliolate 15
2a. Herbs 3
b. Woody plants, trees or shrubby 4
3a. Flowers small, with epicalyx, without petals, with 1-5 stamens. Achenes enclosed
in hard hypanthium Alchemilla (p. 301)
b. Rowers usually showy, without epicalyx, petals only in few species absent, stamens
numerous. Drupes cohering and forming a collective fruit Rubus (p. 247)
4a. Ovary or ovaries superior, not connate with the hypanthium 5
b. Ovary inferior or semi-inferior, carpels connate with the hypanthium, either entirely
or at least their lower half 8
5a. Flower with 1 pistil (exceptionally 2) 6
b. Flower with more than 2 pistils • • • 7
6a. Small shrubs. Leaves 3-lobed. Glands on the hypanthium outside, elongating after
anthesis. Fruits dehiscent, with several seeds Neillia (p. 245)
b. Trees or shrubs. Leaves not lobed. Hypanthium without long-stalked glands. Fruit
a drupe with 1 stone Prunus (p. 319)
7a. Plants climbing, straggling, or creeping, rarely erect, usually armed. Stipules pres-
ent. Fruits cohering as a collective, each of the juicy drupes with one stone
Rubus (p. 247)
b. Erect shrubs, not armed. Stipules absent. Fruit a dehiscent follicle with several seeds
(cultivated) Spiraea (p. 244)
8a. Simple racemes 9
b. Racemes branched and compound 10
9a. Styles free. Fruits with stone-cells in mesocarp (cultivated) Pyrus (p. 317)
b. Styles connate at base. Fruits without stone-cells (cultivated) .... Malus (p. 310)
10a. Plants with thorns (cultivated) Pyracantha (p. 316)
b. Plants without thorns 11
1 1 a. Upper rim of hypanthium and sepals persistent on the fruit. Ovary semi-inferior 12
b. Sepals and upper rim of hypanthium deciduous after flowering, leaving a scar on
top of the fruit. Ovary inferior 14
12a. Main nerves terminating in the margin Eriobotrya (p. 308)
b. Main side-nerves not terminating in the margin 13
13a. Rower with 2 carpels, in their basal half adnate to the hypanthium, but free from
each other (cultivated) Cotoneaster (p. 308)
b. Flower with up to 5 carpels, connate with each other and with the hypanthium
Photinia (p. 3 12)
14a. Evergreen. Ovary 2-celled. Fruits 1-, rarely 2-seeded, seeds thick, cotyledons thick
Rhaphiolepis (p. 317)
b. Deciduous. Ovary usually 3-celled (2-4). Fruits with usually 2 seeds per cell, seeds
flat, cotyledons flat Micromeles (p. 310)
244 Flora Malesiana ser. I, Vol. 11 (2) (1993)
15a. Herbs 16
b. Plants with woody branches 20
16a. Flowers in globular heads, without petals 17
b. Flowers not in heads. Petals present 18
17a. Sepals imbricate. Male and bisexual flowers with 10-30 stamens (cultivated)
Sanguisorba (p. 301)
b. Sepals valvate. Flowers bisexual, with 2 or 3 stamens Acaena (p. 297)
1 8a. Epicalyx absent. Pistils usually 2, remaining enclosed in the hypanthium which bears
many erect spines under the calyx Agrimonia (p. 299)
b. Epicalyx present. Pistils on a more or less elevated torus, not completely enclosed
by the spineless hypanthium 19
19a. Petals yellow Potentilla (p. 286)
b. Petals white (cultivated) Fragaria (p. 285)
20a. Achenes hidden in the hollowed, fleshy hypanthium Rosa (p. 303)
b. Drupes forming a collective fruit, on an elevated torus Rubus (p. 247)
TRIBUS SPIRAEEAE
Mostly woody plants with simple leaves. Stipules absent. Epicalyx absent. Pistils mostly
several, superior, free, not entirely enclosed in hypanthium. Ovules 2 or more, pendu-
lous. Follicles, x = 9.
SPIRAEA
Spiraea L., Sp. PL (1753) 489. — Type species: Spiraea salicifolia L.
Unarmed shrubs. Leaves simple. Stipules absent. Flowers in terminal or axillary um-
bels, corymbs, or panicles. Hypanthium campanulate to turbinate, upper part of disc free,
erect, fleshy. Sepals valvate or slightly imbricate. Petals white or pink. Stamens 15 -many.
Pistils 1-8, often 5; ovaries 1-locular; style (sub)terminal; ovules 2 -several. Follicles
dry, dehiscing ventrally and later at the top also dorsally. Seed with membranous testa,
endosperm thin or absent.
Distribution — About 100 species, North temperate. Many species cultivated as orna-
mentals.
Spiraea cantoniensis Lour., Fl. Cochinchin. Distribution - Southeast China (Guangdong to
1 (1790) 322; Backer & Bakh. f., Fl. Java 1 Zhejiang). The species is widely cultivated, often
(1964) 511. with double flowers. In Malesia cultivated, among
„, , ., L . _ , . . . others in Java (see also Backer & Bakh. f., I.e.,
Shrub with thin, flexuous, glabrous branches. . , . . _ _ . man
j.uul; wiu u , , 6 where also two other species of Spiraea are men-
Petwle 0.5-1 cm. Leaves oblong, (bi)serrate, 3-7
by 1-3 cm, glabrous, ± glaucous underneath. Co-
rymbs axillary, shortly peduncled, with c. 10-20 Run wild from cultivation near Edie Creek,
flowers, pedicels 7-15 mm. Margin of disc lobed. Morobe Prov., Papua New Guinea {Hartley 11673,
Stamens in normal flowers 20-24. Pistils (3-)5. collected in 1963).
Kalkman — Rosaceae
245
Tribus Neillieae
Woody plants with simple leaves. Stipules free, on twig. Epicalyx absent. Usually 1 pistil,
superior, enclosed by hypanthium. Ovules 2 or more, mostly pendulous. Follicles, x = 9.
NEILLIA
Neillia D. Don, Prod. Fl. Nepal. (Febr. 1825) 228; DC, Prod. 2 (Nov. 1825) 546. — Type
species: Neillia thyrsiflora D. Don.
Adenilema Blume, Bijdr. (1826) 1120; Hassk., Cat. Hon. Bog. (1844) 170 (' Adenilem-
ma'). — Type species: Aadenilema fallax Blume.
Unarmed shrubs, erect, scandent, or creeping. Leaves simple, lobed and dentate. Stip-
ules free. Flowers in terminal racemes or panicles, 5-merous, bisexual. Hypanthium cam-
panulate to tubular, often with stalked glands outside. Sepals imbricate, acuminate. Petals
(sub)orbicular, white. Stamens 6-40. Pistil 1, rarely 2, free; ovary superior, included in
hypanthium, 1-locular, style terminal, with peltate stigma; ovules 2-13, biseriately placed
on the ventral placenta. Fruit a dry follicle, protruding from the enlarged hypanthium. Seeds
several, with hard and shining testa; endosperm rather plentiful.
Distribution — About 12 species, from the Himalayan region through China to Korea
and to Vietnam, one species also in Malesia: Sumatra and Java. — Fig. L
Neillia thyrsiflora D.Don, Prod. Fl. Nepal.
(1825) 228; Miq., Fl. Ind. Bat. I, 1 (1855) 391;
Steenis, Bull. Jard. Bot. Buitenzorg III, 13
(1934) 241; Backer & Bakh. f., Fl. Java 1 (1964)
510; Steenis, Mount. Fl. Java (1972) pi. 44-4.
— Types: Wallich 698, Hamilton s.n.
Adenilema fallax Blume, Bijdr. (1826) 1121.—
Neillia fallax (Blume) Blume, Mel. BoL 1 (1855)
6 (see Taxon 35, 1986, 272). — Type: Blume
477, Mt Gede.
Rubus schefferi Focke, Abh. Naturw. Ver. Bremen
8 (1884) 472. — Type: Scheffer s.n., Mt Pan-
gerango.
Neillia lunkinensis Vidal, Not. SysL 13 (1948) 292.
— Neillia thyrsiflora D.Don var. tunkinensis
(Vidal) Vidal, Adansonia 3 (1963) 153. — Type:
Poilane 26671.
Small shrubs, erect, scandent, or creeping.
Branches glabrous, ultimate ones often zigzag.
Leaves ovate, 3-lobed, 4-12 by 2.5-7. 5(-ll) cm,
base cordate to subtruncate, serrate, 3 main nerves,
the middle one with 5-7 side-nerves, nerves and
veins terminating in marginal teeth, glabrous to
sparsely hairy. Petiole 0.5-2 cm. Stipules mem-
branous, up to 12 by 6.5 mm, ± serrate. Inflores-
cence a bracteate compound raceme, the lower of
the up to 8 side-branches in the axils of the upper
leaves, up to 12(-20) cm long, pedicels up to 2
mm. Flowers erect. Hypanthium widely campanu-
late, 1.5-3 mm high, growing after anthesis,
hairy to glabrous outside, always with scattered
stalked glands, the stalks growing to 6 mm after
anthesis. Sepals ± spreading in anthesis, 3-5 by
1-2 mm including die up to 2(-3) mm long
acumen, persistent, indumentum outside as hypan-
thium and often with occasional glands. Petals
suborbicular to ovate, 1.5-3.5 by 1.5-3 mm,
not or shortly clawed, with ciliate margin, early
caducous. Stamens 7-21, filaments up to 1.2 mm,
anthers c. 0.5 mm long. Pistil usually 1, ovary
ovoid, 1.5-2.5 mm long, style up to 2 mm,
ovules 7-10. Follicle 6-9 by 2.5-3.5 mm, not
including the persistent style remnant. Seeds sev-
eral, up to 2 mm long. - Fig. 1.
Notes - Neillia fallax has usually been consid-
ered as conspecific with N. thyrsiflora, but Vidal
(1963) separated the two on a specific level men-
tioning as distinguishing characters: shape of in-
florescence, number of stamens and ovules, indu-
mentum on leaves, inflorescences, and pistils. In
all but one of these characters, however, there is an
unbroken continuity and the only possible way to
break up the complex into recognizable and defin-
246
Flora Malesiana ser. I, Vol. 11 (2) (1993)
able sub-units is to make use of the indumentum
on raceme axis and pedicels. There is no correla-
tion with other characters and the groups are not
geographically separated. In view of this the two
groups do not seem worthy of specific rank.
Neillia rubiflora, a continental- Asian species, is
very closely related but may remain separated, i.a.
because of the larger number of stamens (20-40).
a. var. fallax (Blume) Kalkman, Blumea 37
(1993) 377. — Adenilema fallax Blume, I.e. —
Neillia fallax (Blume) Blume, I.e. — ? Rubus
schefferi Focke, I.e.
Inflorescences glabrous, the pedicels sometimes
with few short hairs. Ovary glabrous or with few
long hairs only.
1 cm
Fig. 1. Neillia thyrsiflora D. Don. a. Flowering branch; b. flower; c. hypanthium with fruits (Schiffner
2014).
Kalkman
Rosaceae
247
Distribution - Malesia: N & W Sumatra, W Java
Habitat - Forest margins and similar open places,
c. 2100-3000 m altitude.
Note - The Sumatran specimens differ from the
Javanese ones in having less hairy leaves and flow-
ers and more stamens (10-21, in Java 7-10).
b. var. thyrsiflora
I.e.
Neillia tunkinensis Vidal,
Inflorescences, incl. pedicels, hairy. Ovary long-
hairy all over or at least with long hairs near suture
and base, rarely (in part of the Sumatra specimens)
glabrous.
Distribution - Continental Asia (Himalayan re-
gion, Assam, China, Vietnam); in Malesia: W Su-
matra and W Java
Habitat - Montane thickets and forest-edges, be-
tween c. 1500 and c. 2500 m altitude.
TRIBUS Rubeae
Woody or herbaceous plants, with simple, pinnate or palmate leaves. Stipules free, on
petiole or twig. Epicalyx absent. Petals rarely reduced in number, flowers rarely uni-
sexual. Pistils many, free, superior, on elevated torus. Ovules 2, pendulous. Drupes.
x = 7. Only genus: Rubus.
RUBUS
Rubus L., Sp. PI. (1753) 492; Focke, Bibl. Bot. 72 (1910-11) 1-223; ibid. 83 (1914)
1-274; Zandee & Kalkman, Blumea 27 (1981) 75; Kalkman, Blumea 29 (1984) 319;
ibid. 32 (1987) 323. — Type species: Rubus fruticosus L. See Taxon 41 (1992) 573.
Shrubs (see under Morphology), usually climbing, straggling or creeping, rarely erect,
only few species herbaceous. Twigs and other parts nearly always with prickles. Leaves
compound (pinnately or palmately structured) or simple, then usually incised. Stipules
free, on the base of the petiole or at the junction of twig and petiole, persistent or fugaci-
ous, rarely absent. Inflorescences terminal, elaborately branched with the lowermost
branches often in the axils of the upper leaves, or little or not branched and in axillary
bundles, or (rarely) strongly reduced and flowers (sub) solitary. Flowers 5-merous, most-
ly bisexual, rarely unisexual and the plants ± dioecious. Sepals imbricate, often unequal,
outer margins often lobed. Petals normally longer than sepals, rarely partly or entirely ab-
sent, white, less commonly cream-coloured, pink, purplish, or red. Stamens many. Pis-
tils many, rarely few, free, on a mostly elevated torus; ovaries 1-locular, style terminal,
stigma capitate or bifid: ovules 2, only 1 developing. Fruits cohering and falling as a col-
lective fruit with or without the torus, or (rarely) coming loose individually, drupes with
usually a juicy or fleshy mesocarp and a hard and rugose endocarp. Seed with thin testa.
— Figs. 2-7.
Distribution — Genus with some hundreds of species, apart from the microspecies in
the apogamous R. fruticosus/ R. caesius complex (according to Weber, Phan. Mon. 7,
1972, there are at least 5000 of those in Europe; see also under Taxonomy). Subcosmo-
politan. In Malesia c. 50 species. New Guinea (17) and the Philippines (c. 17) being rich-
est in species, followed by Java and Sumatra. In the area New Guinea is the only (small)
centre of endemism with 12 endemic species, the other islands have very few endemics or
none at all.
248 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Habitat — Mainly montane with an altitudinal range between 1000 and 3000 m, about
a dozen species normally (also) under 1000 m, only three going down to sea-level, also
a dozen species up into the alpine zone above 3000 m. Highest record is 4340 m for R.
archboldianus (Mt Wilhelm, Papua New Guinea).
Ecology — Most species are light-loving and are restricted to more or less open places,
either natural or anthropogenic. There does not seem to be evidence for suspecting apo-
gamy in any of the Malesian species.
Taxonomy — Focke (1910-11) recognized twelve subgenera but made clear in his
text (p. 6) that he considered the smaller subgenera to be offshoots from the three larger
ones, that represent the main subdivision of the genus: subgenus Malachobatus (centred in
SE Asia), Idaeobatus (centred in E Asia), and Rubus (centred in S America). Nevertheless
he put the large groups and the small offshoots on the same taxonomical level what from a
phylogenetical point of view would not be the most acceptable classification. In the present
survey also the subgenus Micranthobatus is recognized although it may not be a holophyl-
etic group (see Kalkman, 1987). The subgenus Chamaebatus has been maintained with
the same misgivings.
The genus Rubus has a bad reputation among taxonomists, undeserved since the prob-
lems are only caused by one Northern Hemisphere offshoot of the subgenus Rubus, the
so-called 'Moriferi' or R.fruticosuslR. caesius complex. The complex is taxonomically
unsolvable, like others of its kind, because of the facultative apogamy and easy hybridi-
zation with stable progeny. It is possible to find the same 'taxa' year after year, to describe
them, and to recognize differences with other, neighbouring 'taxa'. Over a large area, how-
ever, it is impossible to reach a hierarchic classification with more or less equivalent taxa.
Although 'batologists' admit that their taxa are not comparable, they nevertheless try to
classify them in the common scientific classification, and with predictably poor results. As
one recent example, H.E. Weber (Die Gattung Rubus in NW. Europa, Phan. Monogr. 7,
1972) subdivides the subgenus Rubus in three sections (restricted to the NW European
species). Of the three sections the Eufruticosi is the most important one, the two others
accommodate, respectively, the dewberry (R. caesius) and the hybrids of the latter with
Eufruticosi species. In the section Eufruticosi one of the two subsections contains the
species that some authors consider to be hybrids with R. idaeus (belonging to subgenus
Idaeobatus), the raspberry. The other subsection is divided into eleven series that are
poorly recognizable and definable. As Weber himself says, in placing individual species in
the series there is much room for the individual discretion of different authors.
The rest of the genus, however, behaves perfectly normally: there is a majority of well-
recognizable, clear-cut species, several difficult cases of specific delimitation, and a small
number of complexes like R. moluccanus that are obviously engaged in active speciation,
possibly linked to an enlargement of their habitats by human interference.
Morphology — European blackberry plants (R.fruticosus complex) usually make long
vegetative shoots, called primocanes or turios, during the summer season. After their first
winter a number of shorter mixed shoots appear in the leaf-axils of the primocanes. These
mixed shoots, floricanes, are of determinate growth and terminate in an inflorescence.
After fruiting, in autumn and winter, the floricanes die back and so does a larger or smal-
ler part of the primocanes. New primocanes will, next spring, appear axillary on the lower
Kalkman — Rosaceae 249
nodes of still living parts of the old primocanes, from the subterranean parts, and from
places where overhanging primocanes have rooted. These plants are not really shrubs,
therefore. See Weber, I.e. (1972: fig.2).
Part of the tropical Rubus species may have the same kind of differentiation and peri-
odicity in the branches but from herbarium specimens reconstruction is impossible. Field
observations in the wild and in tropical botanic gardens are needed. In some cases it is
obvious from the herbarium that there are two kinds of branches: long and stout ones with
large leaves in whose axils shorter branches develop, terminating in an inflorescence. Con-
tinuation of growth of these shorter branches, if it occurs, must be sympodial.
Uses — The plants contain tannins, which may be responsible for their use in cases
of diarrhoea and throat-troubles. These uses are reported for tea made from leaves of the
European Rubus fruticosus and also for R. moluccanus and some other Malesian species.
The fruits of all species are edible, but not all are as juicy and tasty as the well-known
American and European blackberries (R. fruticosus I caesius complex), the raspberry (R.
idaeus), loganberry (R. x loganobaccus), or Japanese wine-berry (R. phoenicolasius).
The fruits of these and other species can be eaten raw as table-fruit or made into preserves,
jams, jellies and fruit juices. A few Malesian species could be promising for these pur-
poses, see Kalkman in in E.W.M. Verheij & R.E. Coronel (eds.), Edible fruits and nuts,
Plant Res. SE Asia (PROSEA Handbook) 2 (1991) 277-278 & 355.
KEY TO THE SPECIES
la. Leaves simple 2
b . Leaves compound 24
2a. Underside of leaves with a closed felt of thin, curly hairs, apart from straight, stouter
hairs on nerves and veins 3
b. Not a closed felt on underside of the leaves, leaf surface visible between the hairs 12
3a. Stipules and bracts entire or with few minute teeth 4
b. Stipules and bracts distinctly toothed to deeply incised 5
4a. Inflorescence richly and widely branched. Sepals at anthesis 5-6 mm long
35. R. luzoniensis
b. Inflorescence a compact, few-flowered raceme. Sepals c. 12 mm long
39. R. perfulvus
5a. Flowerbuds globular 6
b. Flowerbuds ovoid, pointed 7
6a. Leaves ± ovate, at least 1.4 times as long as wide, not or very shallowly lobed. Dioe-
cious 31. R. elongatus
b. Leaves suborbicular to broadly ovate in outline, ± as long as wide, deeply 5- to 7-
lobed. Flowers bisexual 26. R. alceifolius
7a. Inflorescence a simple, compact raceme 41. R. rolfei
b. Inflorescence elaborately branched 8
8a. Leaves bullate above, surface distinctly raised between the veins 9
b. Leaves flat above, nerves (and veins) sometimes slightly impressed but leaf surface
between the veins not distinctly raised 1 1
250 Flora Malesiana ser. I, Vol. 11 (2) (1993)
9a. Outer sepals shortly toothed, teeth up to 1.5 mm long. Anthers glabrous
29. R. chrysophyllus
b. Outer sepals with longer lobes or teeth. Anthers hairy on top 10
10a. Outer sepals with 5 or more lobes on each side, lobes up to 7 mm long
33. R. heterosepalus
b. Outer sepals with 1-3 teeth on each side, teeth up to 3 mm long 36. R. malvaceus
11a. Leaves (broadly) ovate, length/width up to 2(-2.5), not or shallowly lobed, if
distinctly lobed (var. angulosus), the basal lobes overlapping. Stipules early falling,
4-12 mm wide, on each side with 4-10 lobes. Ovaries and fruits glabrous
38. R. moluccanus
b. Leaves broadly ovate, length/width less than 1.5, distinctly 3-lobed, base cordate
but lobes not touching. Stipules rather persistent, up to 2 mm wide, with 2 or 3 lobes
on each side. Ovaries and fruits glabrous or hairy 34. R. keleterios
12a. Creeping plants with solitary, terminal flowers, rarely 1 or 2 axillary flowers under
the terminal one. Leaves reniform, often wider than long. Hypanthium and outside
of sepals with needle-shaped prickles 13
b. Erect, climbing, or straggling shrubs with flowers in inflorescences. Leaves usually
distinctly longer than wide. Hypanthium and sepals unarmed 14
13a. Stipules serrate to dentate 45. R. calycinus
b. Stipules deeply digitately divided 46. R. pectinellus
14a. Anthers basifixed 15
b. Anthers dorsifixed or dorso-versatile 16
15a. Underside of leaves with scattered hairs only 42. R. smithii
b. Underside of leaves densely hairy 27. R. beccarii
16a. Ovaries hairy 17
b. Ovaries glabrous, on hairy or glabrous torus 19
17a. Leaves distinctly cordate at base 44. R. sundaicus
b. Leaves with rounded, subtruncate or very shallowly cordate base 18
18a. Leaves less than 1.5 times as long as wide, petiole 1.5 cm or longer, nerves termi-
nating in the margin. Flowers unisexual (?) 30. R. cumingii
b. Leaves normally more than 1.5 times as long as wide, petiole rarely longer than 1 cm,
nerves not reaching the margin. Flowers bisexual 40. R. pyrifolius
19a. Petals 5 20
b. Petals or 1 22
20a. Leaves suborbicular to broadly ovate in outline, about as long as wide, deeply 5-7-
lobed. Stipules and bracts divided into thin thead-like lobes . 26. R. alceifolius
b. Leaves longer than wide, not or hardly lobed. Stipules and bracts toothed or dis-
sected but not with thin, thread-like lobes 21
21a. Inflorescences laxly paniculate, up to 35 cm long, compound racemes with up to 20
racemes of each up to 30 flowers 28. R. benguetensis
b. Inflorescences rarely longer than 10 cm, racemes of 3- to 1 -flowered cymes, the up-
per ones often congested 32. R. glomeratus
22a. Flowers large, sepals 10-12 mm long 43. R. sorsogonensis
b. Flowers smaller, sepals 6-8 mm long 23
Kalkman — Rosaceae 251
23a. Leaves 1-1.3 times as long as wide. Glandular hairs on outside of hypanthium and
sepals, outer sepals on each margin with 3-6 distinct teeth of up to 1 mm long
37. R. mearnsii
b. Leaves 1.4 or more times as long as wide. Outside of hypanthium and sepals with-
out glandular hairs, outer sepals entire or with 1 or 2 minute teeth on each margin
28. R. benguetensis
24a. Leaves pinnate with more than 3 leaflets, or bipinnate 25
b. Leaves 3-foliolate or pedately/palmately 5-foliolate 32
25a. Needle-shaped spines on stems, leaves and also on hypanthium 1 to 5 spines alter-
nating with the sepals 26
b. Prickles not needle-shaped 28
26a. Leaves bipinnate, 3-pinnate at base, simply pinnate at apex
20. R. montis-wilhelmi
b . Leaves simply pinnate 27
27a. Leaflets 1-4 by 0.5-2 cm, with 7— 12(— 18) teeth on each side, upper surface gla-
brous or hairy between the nerves, lower surface glabrous or hairy on midrib and
veins 14. R. ferdinandi-muelleri
b. Leaflets 0.5-1.5 by 0.5-1 cm, with 4-6(-7) teeth on each side, both surfaces gla-
brous or almost so 23. R. papuanus
28a. Stems and leaves with up to 5 mm long gland-tipped hairs . 25. R. sumatranus
b. No long gland-tipped hairs present 29
29a. Leaves almost glabrous 30
b. Leaves hairy, at least on underside 31
30a. Leaflets biserrate, with long-tapering apex. Pistils down to the base of the torus,
ovaries with shortly stalked glands 11. R. chrysogaeus
b. Leaflets serrate, apex acute to long-pointed. Base of torus without pistils. Ovaries
without glands 15. R. fraxinifolius
31a. Leaflets below with a dense, woolly, silvery-white felt all over and with straight
hairs on the nerves. Flowers small: sepals 4-7 mm long, petals 3.5-5 mm long,
pink. Ovaries and fruits densely hairy 22. R. niveus
b. Leaflets below long-hairy but without woolly felt. Rowers large: sepals 7-15 mm
or longer, petals 8-17 mm long, white. Ovaries with stalked glands and apically
with some hairs 24. R. rosifolius
32a. Leaves 5-foliolate, the uppermost ones and/or the first leaves on a branch sometimes
only 3-foliolate 33
b. Leaves 3-foliolate, the upper ones sometimes unifoliolate 37
33a. Leaflets with 18 or (many) more pairs of nerves. Unarmed or prickles very rare 34
b. Leaflets with 6-17 pairs of nerves. Armed 35
34a. Normally 30-40 pairs of nerves. Stipules 2-4 cm long. Sepals sericeous outside,
the covered parts woolly. Petals 4-5 mm long, distinctly shorter than sepals
16. R. Iineatus
b. Normally 16-26 pairs of nerves. Stipules 0.5-1 cm long. Sepals with few hairs
outside, covered parts woolly. Petals 9-12 mm long, ± as long as sepals
21. R. neo-ebudicus
252 Flora Malesiana ser. I, Vol. 11 (2) (1993)
35a. Prickles rather few. Stipules long persistent, 4-12 mm long. Inflorescences mono-
chasial or dichasial 8. R. alpestris
b. Prickles rather many. Stipules absent or fugacious, 3-7 mm long. Inflorescences
racemose, racemes solitary or in bundles 36
36a. Terminal leaflets 4.5-12 by 2-7 cm, with 9-14 pairs of nerves. Racemes with 10-
25 flowers, solitary or in bundles. Filaments glabrous. Ovaries glabrous or hairy,
not glandular 5. R. royenii
b. Terminal leaflets 3-5 by 2-3 cm, with 6-8 pairs of nerves. Racemes with 2-7
flowers, solitary. Filaments long-hairy. Ovaries hairy and dorsally with many yel-
low glands 4. R. novoguineensis
37a. Stems unarmed or almost so 38
b. Stems armed 40
38a. Leaflets with more than 16 pairs of nerves 21. R. neo-ebudicus
b. Leaflets with fewer than 10 pairs of nerves 39
39a. Inflorescence a large, terminal thyrse with up to 20 rich-flowered laterals under the
terminal flower 1. R. dementis
b. Inflorescence much poorer, with under the terminal flower only 1-4 axillary cymes
of up to 4 flowers 18. R. lowii
40a. Hypanthium outside with up to 5 mm long bristles or spine-like hairs, otherwise
hairy or not 41
b. Hypanthium outside without bristles or spines, with or without short hairs and/or
few short prickles 44
41a. Stipules deeply divided. Petals red. Ovaries hairy 9. R. archboldianus
b. Stipules entire, serrate, or with some short teeth. Petals white 42
42a. Hypanthium outside with straight prickles and with capitate spines, otherwise gla-
brous. Ovaries glabrous. Fruits red 12. R. copelandii
b. Hypanthium outside hairy and with straight bristles 43
43a. Stipules 10-18 by 3-15 mm. Inflorescences poor, usually less than 8 flowers. Ova-
ries glabrous. Fruits orange to red 17. R. lorentzianus
b. Stipules up to 10 by 0.5 mm. Inflorescences rich, up to 150 flowers or more. Ova-
ries hairy. Fruits yellow to orange 13. R. ellipticus
44a. Flowers solitary and terminal or in poor racemes of 2-5 flowers 45
b. Inflorescences richer, thyrsoid or racemose 47
45a. Underside of leaflets woolly all over 3. R. megacarpus
b. Underside of leaflets glabrous or with hairs on midrib and larger nerves only . . 47
46a. Ovaries long-hairy 19. R. macgregorii
b. Ovaries glabrous 18. R. lowii
47a. Under the terminal flower up to 6 lateral cymes or dichasia in the axils of bracts or
leaves. Flowers bisexual 48
b. Axillary racemes, almost always simple, rarely partly with cymes instead of flowers,
racemes solitary or in bundles of up to 4. Flowers (always?) unisexual 50
48a. Stipules deeply divided 10. R. banghamii
b. Stipules entire or with 1 or 2 small teeth 49
Kalkman — Rosaceae 253
49a. Petals longer than sepals, early falling. Stamens and pistils both more than 150
7. R. acuminatissimus
b. Petals shorter than sepals, long-persistent Stamens up to 45, pistils up to 25
18. R. lowii
50a. Terminal leaflets 3-10 by 2-7 cm. Flowers small: sepals 2.5-4 mm, petals 4-8
mm long 2. R. diclinis
b. Terminal leaflets 10-15 by 8-12 cm. Rowers larger: sepals 6-7 mm, petals 10-17
mm long 6. R. trigonus
Subgenus Micranthobatus
Rubus subg. Micranthobatus (Fritsch) Kalkman, Blumea 32 (1987) 324. — Rubus sect.
Micranthobatus Fritsch, Osterr. Bot. Zeitschr. 36 (1886) 259.
Leaves palmate, rarely unifoliolate. Leaflets pinninerved, nerves terminating in the
margin. Stipules absent or 1 or 2 on the base of the petiole. Inflorescences terminal thyrses
or axillary bundles of (1— )2— 5 racemes. Flowers uni- or bisexual. Hypanthium saucer-
shaped, sometimes with prickles outside. Sepals (sub)equal, entire. Fruits falling as a co-
herent collective from the torus, rarely falling separately (?).
Distribution — Twelve species with an East Gondwanan type area: Australia, New
Guinea, New Zealand, Celebes, Borneo, Philippines, NE India, Madagascar. In Malesia
6 species.
1. Rubus dementis Merr., Philipp. J. Sc, BoL as are the rachises. Flowers uni- or bisexual, plants
3 (1908) 139; Elmer, Leafl. Philipp. Bot. 2 probably polygamo-dioecious. Hypanthium saucer-
(1908) 458. — Rubus lucens Focke var. clemen- shaped, 3.5-5 mm across, woolly outside. Sepals
tis (Merr.) Focke, Bibl. Bot. 72 (1911) 213; triangular to ovate, 3.5-5 by 3-4.5 mm, obtuse,
Merr., Enum. Philipp. Flow. PI. 2 (1923) 228. apiculate to acuminate, densely short-woolly out-
— Type: Clemens 740, Mindanao. side. Petals obovate to elliptic, 5.5-9 by 4-6
mm, white. Stamens 70-160, filaments up to 2.5
Climbing or scrambling shrubs. Stems up to mm, anthers 1-1.5 mm long, staminodes in female
20 m long, unarmed or with few, short, curved flowers 1-2 mm, including minute anther rudi-
prickles, short-hairy and with minute, stalked ment. Pistils 30-70, ovaries glabrous or with
glands. Leaves 3-foliolate, petiole 5-9 cm long. long hairs dorsally near apex, on elevated, hairy
Stipules on the peuole, up to 5 mm above its torus, style 2-3 mm long, sometimes hairs at
base, linear, up to 7 by 1 mm, rather persistent. base, pistillodes in male flowers 2 mm including
Leaflets ovate to elliptic, terminal ones 8-13.5 by style. Collective fruits globular, 1-1.5 cm, sepals
4.5-9 cm, lateral ones usually slightly smaller, closing and enlarging after anthesis. Fruits densely
base rounded, margin shallowly serrate in upper packed, 2.5-3.5 mm by 1.5-2 mm when dry,
part, apex acuminate, nervation pinnate with 7-9 exocarp sometimes still hairy, red to orange-red,
pairs of nerves, often near the end with one strong mesocarp juicy.
acropetalous tertiary vein, venation transverse, Distribution - Northern part of Sumatra, Borneo,
sparsely short-hairy and with minute glands. In- Mindanao, Celebes.
florescence terminal, up to 55 cm long, a hanging Habitat - In forest along rivers and brooks, also
thyrse with up to 20 laterals in the axils of bracts in open places in forest and shrubland, altitude
or leaves, the lower laterals up to 25 cm long. 150-1300(-1800) m.
Bracts linear, up to 5 mm long. Pedicels up to 5 Note - The disposition of uni- and bisexual flow-
mm long, growing to 10 mm, densely short-hairy ers is incompletely known, see Kalkman (1987).
254
Flora Malesiana ser. I, Vol. 11 (2) (1993)
2. Rubus diclinis F. Muell., Trans. Roy. Soc.
Vict. 1 (1889) 5; P. van Royen, Phan. Mon. 2
(1969) 69, only var. diclinis; Alpine Fl. New
Guinea 4 (1983) 2481, idem. — Types: Mac-
Gregor s.n., Mt Knutsford, Mt Musgrave.
Rubus tsiri P. van Royen, Phan. Mon. 2 (1969)
77, excl. most specimens cited; Alpine Fl. New
Guinea 4 (1983) 2486, idem. — Rubus para-
doxus Ridley, Trans. Linn. Soc. Lond. II, BoL
9 (1916) 36, nom. illeg., non S.Moore (1878).
— Types: Kloss specimens, Mt Jaya (Carstensz).
Climbing or scrambling shrubs. Stems up to
5(-18?) m long, densely patendy hairy, sometimes
with shortly stalked glands, prickles rather many,
curved, small. Leaves 3-foliolate, petiole 1.5-5.5
cm long. Stipules very rarely present, on the peti-
ole, up to 4 by 0.5 mm. Leaflets elliptic to ovate,
terminal ones 3-10 by 2-7 cm, lateral ones smal-
ler, base rounded to shallowly cordate, margin ser-
rate, apex acute, hard-chartaceous, nervation pinnate
with (6-)8-12 pairs of nerves, venation trans-
verse, scattered hairy above, sparsely to densely
golden to brownish hairy below. Inflorescences
simple racemes, solitary or in bundles of up to 4
in the leaf axils, up to 8 cm long, peduncle 0-2
mm, some empty bracts at base, up to 15 flowers.
Bracts up to 7 by 4 mm. Pedicels 4-10 mm long,
densely hairy and always with small curved prick-
les, as is the rachis. Flowers normally unisexual,
sometimes bisexual (at least appearing so in the
herbarium). Hypanthium saucer-shaped, 2.5-3 mm
across, woolly and with patent hairs outside. Sepals
elliptic to tongue-shaped, 2.5-4 by 1.5-3.5 mm,
with straight hairs outside and woolly on the mar-
gins, pinkish or purplish inside. Petals elliptic to
oblong, 4-8 by (l-)2.5-4.5 mm, obtuse or emar-
ginate, patently hairy inside, white or pale pink.
Stamens 18-40, filaments up to 4.5 mm, anthers
0.5-1 mm long, staminodes in female flowers
small and ± petaloid. Pistils 10-20, ovaries hairy
or glabrous, on elevated, densely hairy torus, style
up to 1.5 mm long, pistillodes in male flowers
minute. Collective fruits up to 1.5 cm when living,
1 cm when dry, sepals spreading. Fruits well sepa-
rated, up to 5(-7) by 4 mm when dry, exocarp hairy
or glabrous, dark red to black, mesocarp rather
thick and fleshy when living, rather thin when dry.
Distribution - New Guinea.
Habitat - Montane forest, clearings, forest edges,
secondary forest, shrubland, altitude 1750-3470 m,
rarely lower.
Uses - On one herbarium label the leaves are
reported to be used "in smoking", i.e. as cigarette
wrapper.
3. Rubus megacarpus P. van Royen, Phan.
Mon. 2 (1969) 65; Alpine Fl. New Guinea 4
(1983) 2480. — Type: Brass 30099, Mt Wil-
helm.
Climbing or scrambling shrubs. Stems up to
10 m long, densely hairy, prickles many, short,
curved, some cataphylls at base of lateral shoots.
Leaves 3-foliolate, petiole 1.5-5 cm long. Stip-
ules not always present, on the petiole, filiform to
linear, 4-6 by 0.1-0.5 mm, early falling. Leaf-
lets obovate to elliptic, 3-6 by 2-3 cm, lateral
ones usually relatively narrower than apical one,
shallowly serrate in upper part only, apex rounded
to truncate, with or without apiculus, stiff-coria-
ceous, nervation pinnate with 5-7 pairs of steeply
ascending nerves, venation transverse, glabrous
above, lower surface densely woolly all over and
with appressed straight hairs on midrib and nerves.
Inflorescence a simple raceme, axillary, up to 12
cm long, rachis stiff, with up to 5 flowers, often
reduced to only the terminal flower. Bracts 5-9 by
1.5-3 mm, persistent, also empty ones under the
flowers. Pedicels up to 4 cm long, densely hairy as
is rachis. Flowers uni- or bisexual. Hypanthium
saucer-shaped, 6-10 mm across, densely hairy out-
side. Sepals broadly ovate, 9-12 by 8-11 mm,
inner ones slightly narrower than outer ones, ob-
tuse and apiculate, shortly woolly and with appres-
sed hairs outside, pale purplish. Petals obovate,
13-20 by 8-13 mm, densely long-hairy in basal
half outside, (pinkish or greenish) white. Stamens
70-100, filaments up to 10 mm, with long hairs,
anthers up to 2 mm long, staminodes in female
flowers minute. Pistils c. 150, ovaries densely
hairy, on high, densely hairy torus, style up to 3
mm long, pistillodes in male flowers minute. Col-
lective fruits large, up to 5.5 by 4 cm when living,
most of the pistils developing, sepals appressed to
spreading after anthesis. Fruits closely packed, up
to 9 by 7 mm when dry, exocarp densely whitish-
hairy, purplish, mesocarp still thick when dry,
stone up to 8 by 6 mm. - Fig. 2.
Distribution - New Guinea, only known from
Mt Wilhelm.
Habitat - Alpine shrubland and mossy forest,
altitude 3500-3750 m.
4. Rubus novoguineensis Merr. & Perry, J.
Arnold Arbor. 21 (1940) 183; Kalkman, Blumea
32 (1987) 337. — Rubus diclinis F. Muell.
var. novoguineensis (Merr. & Perry) P. van
Royen, Phan. Mon. 2 (1969) 75; Alpine Fl.
New Guinea 4 (1983) 2484. — Type: Brass
4337, Mt Albert Edward.
Kalkman — Rosaceae
255
Fig. 2. Rubus megacarpus P. van Royen. Fruiting twig (Brass 30099). Photo P. van Royen.
Scrambling or trailing shrubs. Stems up to 2 m
long, densely hairy and with few stalked glands,
glabrescent, prickles rather many, curved, up to 2
mm, several cataphylls at base of laterals. Leaves
palmately 5-foliolate, the first leaves on a branch
sometimes 4- or 3-foliolate, petiole 2.5-5.5 cm
long, petioles and petiolules hairy, with rather
many strongly curved prickles. Stipules maybe not
always present, on the petiole, 3-5 by less than
1 mm, hairy. Leaflets elliptic, terminal ones 3-5
by 2-3 cm, base rounded to slightly cordate, mar-
gin serrate, apex rounded, chartaceous, nervation
pinnate with 6-8 pairs of nerves, venation trans-
verse, densely (semi-)patendy hairy below, glabres-
cent. Inflorescence a simple raceme, axillary, 5-
7.5 cm long, with 2-7 flowers, peduncle up to
2 cm, with some to many prickles. Bracts up to
6 by 2 mm, also empty ones at base of peduncle.
Pedicels 1.5-4 cm long, with prickles. Flowers
unisexual and plants dioecious or polygamodioe-
cious. Hypanthium flat, 3-3.5 mm across, ap-
pressed-hairy and glandular outside, sometimes
with a prickle. Sepals elliptic, 3.5-7 by 3-6 mm,
obtuse, indumentum outside as hypanthium, pur-
plish. Petals elliptic, 5-7 by 3.5-7 mm, rounded
to slightly emarginate at apex, hairy, pale green to
white. Stamens 14-20, filaments up to 2.5 mm,
long-hairy, anthers c. 1 mm long, staminodes in
female flowers minute. Pistils 17-25 in a com-
pact globe, ovaries densely long-hairy and yellow-
glandular on the backside, on a flat hairy torus,
style 0.8-1 mm long. Collective fruits globular,
1-1.5 cm when dry, up to 2.5 cm when living,
sepals spreading. Fruits well separated, 5-7 by
4-4.5 mm, exocarp hairy and dorsally also glan-
dular, brown (?), mesocarp thick.
Distribution - New Guinea, only known from
Central Prov. in Papua New Guinea.
Habitat - Open places in forest and in forest mar-
gins, altitude 2800-3680 m.
256
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Note - Closely related is the Australian Rubus
moorei F. Muell., under which name two species
are hiding. See Kalkman, I.e.: 334, 338.
5. Rubus royenii Kalkman, Blumea 32 (1987)
333. — Rubus tsiri auct. non P. van Royen:
P. van Royen, Phan. Mon. 2 (1969) 77, excl.
type; Alpine Fl. New Guinea 4 (1983) 2486,
excl. type. — Type: Brass 30919, Mt Otto.
Rubus diclinis F. Muell. var. ikilimbu P. van
Royen, Phan. Mon. 2 (1969) 75; Alpine Fl. New
Guinea 4 (1983) 2486. — Rubus royenii Kalk-
man var. ikilimbu (P. van Royen) Kalkman,
I.e.: 336. — Type: see below.
Rubus moorei auct. non F Muell.: Merr. & Perry,
J. Arnold Arbor. 21 (1940) 184, in obs.
Climbing, scrambling, or trailing shrubs.
Stems up to 6(-10) m long, variously hairy,
prickles rather many, 1-2 mm long, straight to
slightly curved, shoots with large cataphylls at
base. Leaves palmately 5-foliolate, occasionally
4- or 3-foliolate, petiole 3-11 cm long, with many
small, curved prickles. Stipules usually absent.
Leaflets elliptic to ± ovate, terminal ones 4.5-12
by 2-7 cm, shallowly cordate to rounded at base,
margin dentate- serrate, apex acute to acuminate,
chartaceous, nervation pinnate with 9-14 pairs of
nerves, rather often forking, venation transverse,
indumentum various. Inflorescences simple ra-
cemes, 1-5 in the leaf axils, 6-15 cm long, pe-
duncle 0-1 cm, with up to 25 flowers. Bracts 3-7
by 1-4 mm, also empty ones at base of peduncle.
Pedicels 4-18 mm long, densely hairy and with
small prickles, as is rachis. Flowers unisexual,
plants probably dioecious. Hypanthium saucer-
shaped, densely hairy outside, sometimes with
prickles. Sepals ± elliptic, obtuse, densely hairy
outside on the marginal parts, pinkish. Petals
elliptic, obtuse to emarginate, long-hairy inside,
white to pink. Stamens glabrous, staminodes in
female flowers minute. Ovaries glabrous or with
long hairs in upper part, on (slightly) elevated,
hairy torus, pistillodes in male flowers minute.
Collective fruits globular, up to 2 cm, probably
late in attaining their final dimensions, sepals
spreading. Fruits dark red to black, with thick
mesocarp.
Distribution - New Guinea, New Britain.
Habitat - Open places and forest, up to 3400 m
altitude, rarely collected below 1200 m.
Note - Related is the Australian Rubus moorei
F Muell., under which name two species are hiding.
See Kalkman, I.e.: 334, 338.
KEY TO THE VARIETIES
la. Long, non-glandular bristles of 3-4(-5) mm
long on stems, petioles and petiolules
a. var. hispidus
b. Bristles absent 2
2a Stalked glands absent. Leaves glabrous to
sparsely hairy on both sides. Flowers small:
sepals 3-4 mm, petals 5-7.5 mm long, sta-
mens 20-40, pistils 10-30 c. var. royenii
b. Stalked glands on stems, petioles, inflores-
cences, and pedicels. Leaves hairy on upper
surface, densely so on midrib and large nerves,
lower surface densely soft-hairy on all nerves
and veins. Flowers larger: sepals 3.5-5 mm,
petals 8-10 mm long, stamens 45-60, pis-
tils c. 50 b. var. ikilimbu
a. var. hispidus Kalkman, Blumea 32 (1987)
336. _ Type: Sayers & Millar NGF 19884, Mt
Wilhelm.
Stems with many, reddish to brown, 3-4(-5)
mm long, straight, non-glandular bristles, other-
wise sparsely hairy to glabrous. Stipules rarely
present, on the petiole, linear, 3-7 mm long. Peti-
oles and petiolules densely hairy and with bristles.
Leaflets on upper surface glabrous, lower surface
sparsely short-hairy on main nerves and sometimes
minutely glandular. Hypanthium 1.5-2.5 mm
across. Sepals 2-3 by 1.8-3 mm. Petals 4.5-6.5
by 2-3 mm, white. Stamens 14-20, filaments up
to 1.5 mm. Pistils 10-20. Collective fruit up to
1.5 cm (living), fruits growing to 6 by 4.5 mm.
Distribution - Papua New Guinea, New Britain.
Habitat - Forest, forest edges, disturbed places,
shrubland, altitude (1800-)2400-3400 m.
Note - One chromosome count 2n = 28 was made
by Borgmann, Zeitschr. f. Bot. 52 (1964) 124, as
Rubus spec, with Borgmann 203 as voucher.
b. var. ikilimbu (P. van Royen) Kalkman, Blu-
mea 32 (1987) 336. — Rubus diclinis F. Muell.
var. ikilimbu P. van Royen, Phan. Mon. 2
(1969) 75. — Type: Hoogland & Pullen 6179,
Upper Wahgi Valley.
Stems, petioles, and petiolules densely soft-
hairy and often with stalked glands, their stalks up
to K-2.5) mm long. Leaflets on upper surface
hairy all over, densely so on main nerves, ± gla-
brescent, on lower surface densely patently soft-
hairy on all nerves and veins. Inflorescence rachis
and pedicels with stalked glands. Hypanthium 3.5-
Kalkman — Rosaceae
257
4.5 mm across, sometimes with stalked glands
outside. Sepals 3.5-5 by 2.5-3. 5(-5) mm. Petals
8—10 by 4-5 mm, pink. Stamens 45-60, fila-
ments up to 2.5 mm. Pistils c. 50. Collective fruit
up to 2 cm (dry), fruits up to 8 by 5.5 mm (dry).
Distribution - Papua New Guinea.
Habitat - Secondary forest and shrubland, alti-
tude 1280-2560 m.
Uses - Stems used for making ropes, leaves are
smoked (Note on Flenley ANU 2071 from Wabag).
c. var. royenii
Stems, petioles, and petiolules sparsely hairy to
glabrous. Leaflets on both surfaces sparsely hairy
on nerves to glabrous. Hypanthium 2.5-3 mm
across. Sepals 3-4 by 2.5-3 mm. Petals 5-7.5
by 2.5-3.5 mm, (pinkish- or cream-)white. Sta-
mens 20-40, filaments c. 2.5 mm. Pistils 10-30.
Collective fruit c. 7 mm, fruits up to 4 by 3.5
mm when dry.
Distribution - New Guinea.
Habitat - Forest margins, secondary forest and
shrubland, streambanks; alt (670-) 1900- 3340 m.
6. Rubus trigonus Kalkman, Blumea 37 (1993)
378. — Rubus cordiformis Kalkman, Blumea
32 (1987) 331, non Weber & Martensen, Son-
derb. Naturwiss. Ver. Hamburg 4 (1981) 100.
— Type: Brass 30932, Mt Wilhelm.
Climbing or scrambling shrubs. Stems up to
6 m long, densely patently hairy and with small,
subsessile glands, prickles rather many, 1-1.5 mm
long, large cataphylls at base of shoots. Leaves 3-
foliolate, petiole 5-12 cm long. Stipules not seen.
Leaflets ovate, terminal ones 10-15 by 8-12 cm,
lateral ones slightly smaller, basal part usually
folded back in herbarium, base cordate, margins
irregularly dentate, apex acuminate, nervation pin-
nate (pedate) with 6-9 pairs of nerves, the lower-
most with some strong basiscopic side-nerves, ve-
nation transverse, patently hairy, nerves on under-
side with glands. Inflorescence very lax, usually a
simple raceme, sometimes partly branched from
the bracteoles, up to 20 cm long, peduncle 0-2
mm long, racemes solitary or 2 or 3 in the axils of
leaves or cataphylls, with up to 10 flowers. Bracts
4-10 mm long. Pedicels 2-5.5 cm long, densely
hairy as is the rachis. Flowers unisexual, plants
probably dioecious. Hypanthium saucer -shaped,
4-5 mm across, densely hairy outside. Sepals el-
liptic, 6-7 by 4-7 mm, obtuse, with patent hairs
and glands outside, the marginal parts woolly.
Petals elliptic, 10-17 by 4-10 mm, long-hairy at
base inside, white or cream-coloured. Stamens 40-
60, filaments up to 7 mm, anthers 1-1.5 mm long,
staminodes in female flowers minute. Pistils c. 60,
ovaries densely hairy, on slightly elevated, hairy
torus, style 1 mm long, pistillodes in male flow-
ers c. 30, minute to small. Collective fruits ellip-
soid, c. 1.2 by 1 cm, sepals spreading under the
ripe fruits. Fruits 4 by 3 mm when dry, exocarp
and mesocarp forming a thin layer when dry, hairy
in upper part, colour unknown.
Distribution - New Guinea (only known from
Papua New Guinea).
Habitat - Shrubland and forest (margins), alti-
tude 1500-3200 m.
Notes - Dr. A. A. van de Beek drew my atten-
tion to the older homonym.
See also under Dubious names, Rubus diclinis
F. Muell. var. papuanus Focke.
Subgenus Ideobatus
Rubus subg. Idaeobatus (Focke) Focke, Bibl. Bot. 72 (1911) 128; Zandee & Kalkman,
Blumea 27 (1981) 79-113. — Rubus sect. Idaeobatus Focke, Abh. Naturw. Ver.
Bremen 4 (1874) 147.
Leaves 3-foliolate, palmately 5-foliolate, or imparipinnate, rarely bipinnate, rarely
1-foliolate (not in Malesia). Leaflets pinninerved, nerves terminating in the margin. Stip-
ules on the basis of the petiole, persistent. Inflorescences thyrsoid, ± elaborate, terminal or
sometimes also lateral. Flowers bisexual. Hypanthium saucer-shaped. Sepals (sub)equal,
usually entire. Fruits cohering, becoming loose from the torus as a whole, endocarp rugose.
Distribution — Many species, distributional centre in Asia, extending to Australia and
the Pacific islands, Africa including Madagascar, islands in the Indian Ocean, few in N
and C America, one species (R. idaeus) in Europe. In Malesia 19 species.
258
Flora Malesiana ser. I, Vol. 11 (2) (1993)
7. Rubus acuminatissimus Hassk., Tijd. NaL
Gesch. Phys. 10 (1843) 146, excl. syn. Rubus
moluccus parvifolius Rumph.; Miq., Fl. Ind.
Bat. I, 1 (1855) 377; Backer & Bakh.f., Fl.
Java 1 (1964) 514. — Type: probably a living
plant in the Bot. Gard. Buitenzorg (Bogor), not
maintained in herbarium.
Rubus podocarpus Kuntze, Rev. Gen. PI. 1 (1891)
223. — Type: Kuntze 5350, Java.
Climbing shrubs, sometimes erect? Stems up
to 3 m long, glabrous, prickles rather many, curved,
stout. Leaves 3-foliolate, usually 1-foliolate near
and in the inflorescence, petiole 1.5-5 cm long.
Stipules linear, 3-7 by up to 1 mm, with some
long hairs. Leaflets elliptic to oblong or ovate-
oblong, terminal one 4-11 by 2-6 cm, lateral
ones 2-9 by 1-4 cm, base rounded, margin ser-
rate, apex acuminate, thinly herbaceous, 6-13
pairs of nerves, upper surface with patent hairs on
main nerves, sometimes appressed hairs between
them, lower surface glabrous except some hairs on
larger nerves. Inflorescence lax, up to 5 cymes be-
low the terminal flower, the cymes 1- to 3 -flowered.
Bracts often leaf-like or 3 -partite. Pedicels up to 5
cm long, glabrous, with prickles. Hypanthium 5-6
mm across, glabrous outside, unarmed or with few
short prickles. Sepals triangular to narrowly ovate,
6-12 by 3-6 mm, growing to 17 mm after an-
thesis, pointed, entire, glabrous outside but woolly
on the covered margins, thick and hard. Petals ob-
ovate, 12-15 by c. 8 mm, falling early, emargi-
nate, with ciliate undulate margin, white. Stamens
150-180, filaments up to 5 mm, anthers c. 1 mm
long. Pistils more than 150, ovaries glabrous, on
elevated, glabrous torus, the lower part without
pistils and stalk-like, style up to 2.5 mm. Collec-
tive fruits (depressed) globose, up to 1.5 cm, sepals
ultimately recurved. Fruits c. 2 by 1 mm (dry),
exocarp orange-red to red, mesocarp a thin mem-
branous layer when dry.
Distribution - Sumatra, W Java.
Habitat - Forest edges and lighter places in for-
est, altitude 1450-2200 m, descending to 700 m
along watercourses [Backer & Bakh. f., Fl. Java 1
(1964)514].
8. Rubus alpestris Blume, Bijdr. (1826) 1108;
Miq., Fl. Ind. Bat. I, 1 (1855) 378; Backer &
Bakh. f., Fl. Java 1 (1964) 514; Steenis, Mount.
Fl. Java (1972) pi. 45-1; Naruhashi & Sato,
Tukar-Menukar 2 (1983) 11. — Type: Blume
407, Java.
Climbing or erect shrubs, up to 4 m. Stems
sparsely hairy when young, with many to few
short glandular hairs, prickles rather few, up to 7
mm, curved to straight. Leaves palmately (some-
times ± pedately) 5-foliolate, upper ones often 3-
foliolate, petiole (l-)2-5 cm long. Stipules linear
to linear-lanceolate, 4-12 by 0.5-2 mm (larger
and wider in Celebes and Moluccas), entire or with
some small teeth, with some glandular hairs, other-
wise ± glabrous. Leaflets oblong, rarely obovate-
oblong, terminal ones (4-)6-14 by 2-4 cm, lat-
eral ones smaller, base acute, margin serrate to
biserrate, apex acuminate to caudate, papyraceous
to pergamentaceous, 9-17 pairs of nerves, upper
surface with few hairs, lower surface hairy. Inflo-
rescences in the axils of the upper 1-3 leaves,
apex of flowering twig usually (?) aborted, dicha-
sial with 1-6 flowers, peduncle up to 5 cm. Bracts
elliptic to lanceolate, up to 12 mm long. Pedicels
up to 3 cm long, sparsely pubescent and with glan-
dular hairs. Hypanthium 6-8 mm across, glabrous
to sparsely hairy outside. Sepals ovate-triangular
to narrowly triangular, 10-15(— 17) by 2.5-6 mm,
entire or outer ones with 1-2 marginal teeth, acu-
minate to long-caudate, acumen up to 6 mm, few
hairs outside but covered margins shortly woolly.
Petals orbicular to elliptic, 6-7 by 5-6 mm, fall-
ing early, with few hairs inside, light green to
white or pink. Stamens c. 50-60, filaments up
to 4 mm, anthers 1-1.5 mm long. Pistils 15-25,
ovaries glabrous, on little elevated to flat, hairy
torus, style up to 8 mm long. Collective fruits
ovoid, c. 1 cm, sepals upright. Fruits up to 4 by
2.5 mm (dry), red, mesocarp moderately thin when
dry.
Distribution - N Thailand, N Vietnam; Malesia:
Sumatra, Borneo, Java, Celebes, Moluccas.
Habitat - Lighter places in forest and shrubland,
altitude 1650-2850(-3000) m.
Uses - Fruits edible, the species even recom-
mended for planting by Koorders (on Koorders
31658).
9. Rubus archboldianus Merr. & Perry, J.
Arnold Arbor. 21 (1940) 180; P. van Royen,
Phan. Mon. 2 (1969) 57; Alpine Fl. New Gui-
nea 4 (1983) 2477. — Type: Brass 4565, Whar-
ton Range.
Climbing or scrambling shrubs. Stems up to
5 m, with long hairs, glabrate, prickles up to 2 mm,
curved, often purplish. Leaves 3-foliolate, upper
ones sometimes simple, petiole 1-4 cm long.
Stipules deeply divided into up to 6 linear lobes,
4-15 mm long, hairy outside. Leaflets elliptic,
ovate-elliptic, or obovate-elliptic, sometimes del-
toid or rhomboid, terminal ones 1.5-12 by 2-6.5
Kalkman — Rosaceae
259
cm, lateral ones up to 5.5 by 4 cm, base usually
acute, margin serrate, apex obtuse to acute or acu-
minate, coriaceous, 5— 10(— 12) pairs of nerves, up-
per surface more or less densely long-hairy, lower
surface with hairs mainly on the nerves. Inflores-
cences with 1 or 2 cymes under the terminal flower,
the cymes 1- or 2-flowered. Bracts stipule-like.
Pedicels up to 4 cm long, hairy and with some
prickles. Hypanthium up to 15 mm across, long-
hairy and with many straight prickles outside,
prickles up to 5 mm. Sepals ovate to triangular,
12-18 by 6-10 mm, growing after anthesis,
caudate, exposed margins with up to 15 long and
slender teeth, indumentum outside as hypanthium,
purplish or reddish. Petals obovate or spathulate,
distinctly clawed, 11-18 by 7-10 mm, early fal-
ling, (orange or pinkish) red. Stamens 50-75, fila-
ments up to 10 mm, anthers up to 1.5 mm long.
Pistils 35-90, ovaries long-hairy in upper part,
on elevated, glabrous torus, style up to 7 mm long,
hairy at base. Collective fruits depressed ovoid, up
to 3 cm. Fruits 4-5 by 2-2.5 mm, exocarp long-
hairy and with a silky shine, dark red, mesocarp
juicy, endocarp dorsally keeled.
Distribution - New Guinea, only known from
the Eastern part
Habitat - In and along edges of different kinds
of mountain forest and in shrubland, altitude 1800-
36O0(-4300) m.
Uses - The fruits are edible and have a pleasant
taste.
Note - According to Borgmann, Zs. f. Bot. 52
(1964) 144 (sub R. spec.3), the species is high-
polyploid with 2n = c. 91.
10. Rubus banghamii Merr., Contr. Arnold
Arbor. 8 (1934) 68, pi. III. — Type: Bangham
1163, Tapanuli.
Semi-scandent shrubs. Stems glabrous, prickles
rather few, curved, up to 5 mm. Leaves 3-foliolate,
upper ones simple and lobed, petiole 1.5-4 cm
long. Stipules deeply 3- to 8-laciniate, lobes up to
12 mm long and up to 0.5 mm wide, glabrous.
Leaflets elliptic, terminal ones 8-10 by 4-5 cm,
lateral ones 6-8 by 2.5-4 cm, base acute, margin
serrate, apex acuminate, herbaceous, 8-12 pairs of
nerves, only short hairs on midrib above, otherwise
glabrous. Inflorescence lax, up to 6 dichasia of up
to 3(-6) flowers below the terminal flower. Bracts
entire or like the stipules. Pedicels up to 6 cm long.
Hypanthium c. 7 mm across (in fruit), glabrous
and unarmed outside. Sepals triangular to triangu-
lar-ovate, 9-17 by 5-8 mm, entire, pointed, gla-
brous outside but woolly on the covered margins.
Petals not seen. Stamens c. 50, filaments up to 7
mm, anthers c. 1 mm long. Pistils c. 35, ovaries gla-
brous, on elevated, densely long-hairy torus, styles
up to 5 mm long. Collective fruits ± ovoid, c. 1.5
cm, sepals spreading. Fruits up to 4 by 2.5 mm,
red, mesocarp a thin membranous layer when dry.
Distribution - Sumatra.
Habitat - Primary forest, 1250-1400 m altitude.
Note - Insufficiently known species.
11. Rubus chrysogaeus P. van Royen, Phan.
Mon. 2 (1969) 42, fig. 8. — Type: Womersley
& van Royen NGF 5919 = van Royen 4334,
Edie Creek.
Sprawling to erect shrubs, up to 2 m. Stems
glabrous, prickles up to 4 mm, slightly curved.
Glands (sub)sessile, pale, on many parts of the
plants. Leaves imparipinnate, with 2 or 3 opposite
pairs of leaflets, often simple in the inflorescence,
up to 17 cm long, petiole 0.8-5 cm long. Stipules
linear, 6-9 by 0.5 mm, entire, ± glabrous. Leaf-
lets ovate-lanceolate to lanceolate, 2-8 by 1-3
cm, base ± rounded, margin biserrate, apex long-
tapering, papyraceous, 9-13 pairs of nerves, both
sides sparsely hairy on main nerves only. Inflores-
cences with up to 4 monochasial or sometimes di-
chasial branches under the terminal flower, alto-
gether up to 12 flowers. Bracts leaf-like to linear.
Pedicels up to 4 cm long, glabrous, with some
small prickles. Hypanthium up to c. 5 mm across,
glabrous outside but with rather many sessile, yel-
low glands. Sepals narrowly triangular, 8-16 by
3-5 mm, incl. the long acumen, indumentum
outside as hypanthium and woolly on the covered
margins. Petals elliptic to obovate or spathulate,
8-12 by 3.5-5 mm, falling early, obtuse, shortly
hairy outside, white. Stamens 70-80, filaments
up to 5 mm, anthers 0.8 mm long. Pistils c. 500,
ovaries (sub)glabrous and with pale glands, on
elevated, hairy torus, style up to 1.5 mm long.
Collective fruits globose to slightly ovoid, up to
1.5 cm, sepals spreading. Fruits up to 1 by 0.8
mm, red, mesocarp only a thin layer when dry.
Distribution - New Guinea, only known from
the Eastern part of Papua New Guinea.
Habitat - Shrubland, forest borders, roadsides
and similar rather open places, also recorded from
grassland and forest, altitude 1200-2600 m.
12. Rubus copelandii Merr., Philipp. J. Sc. 1.
Suppl. 3 (1906) 194 (' copelandii: Elmer. Leafl.
Philipp. Bot. 2 (1908) 457; Merr., Enum.
Philipp. Flow. PI. 2 (1923) 227. — Type: Mer-
rill 4810, Pauai.
260
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Climbing or sprawling shrubs. Stems up to 4 m,
with spine-like capitate hairs, otherwise glabrous,
prickles many, up to 5 mm, straight to curved.
Leaves 3-foliolate, in the inflorescence and at the
base of laterals often simple and lobed, petiole
2.5-6.5 cm long. Stipules ovate to oblong, 7-15
by 2-7 mm, entire or with some teeth, with capi-
tate hairs. Leaflets ovate, terminal ones 4-10 by
3-7 cm, lateral ones slightly smaller, base ± round-
ed, margin (bi)serrate, apex acute to acuminate,
herbaceous, 9-12 pairs of nerves, upper surface
shortly hairy on main nerves and with long appres-
sed hairs between them, lower surface glabrous. In-
florescences lax, up to 3 dichasia of up to 5 flowers
under the terminal flower. Bracts stipule-like or
3-partite. Pedicels up to 6 cm long, with curved
prickles and capitate spines. Hypanlhium c. 5 mm
across, glabrous outside but with straight prickles
and capitate spines. Sepals triangular to ovate, 7-11
by 3-5 mm, entire, acuminate, indumentum out-
side as hypanthium and the covered parts woolly.
Petals broadly elliptic to obovate, 9-12 by 7-9
mm, early falling, rounded, sometimes ciliolate,
white. Stamens 80-100, filaments up to 5 mm,
anthers c. 1 mm long. Pistils over 100, ovaries
glabrous, on elevated, glabrous torus, style up to
2 mm long. Collective fruits ovoid, up to 2 by 1
cm, sepals spreading. Fruits up to 2 by 1.5 mm,
dark red, mesocarp a thin membranous layer when
dry.
Distribution - Luzon.
Habitat - More or less open places in forest,
forest borders, thickets, altitude 1700-2450 m.
13. Rubus ellipticus J.E. Sm. in Rees, Cyclop.
30 (1815) Rubus spec. 16; Elmer, Leafl. Philipp.
Bot. 2 (1908) 456; Merr., Enum. Philipp. Row.
PI. 2 (1923) 227; Backer & Bakh. f., Fl. Java 1
(1964) 515. — Type: Hamilton s.n., Nepal.
Climbing or scrawling shrubs. Stems up to 4 m,
densely woolly when young, with many patent red
bristles (up to 8 mm long and glandular when
young), prickles rather few, straight to slightly
curved, up to 8 mm. Leaves 3-foliolate, the upper
ones sometimes simple, petiole 1-7.5 cm long.
Stipules linear, up to 10 by 0.5 mm, entire, hairy.
Leaflets elliptic to orbicular, sometimes slightly
ovate or obovate, terminal ones 4-9 by 3.5-9 cm,
lateral ones 2-6.5 by 2-6.5 cm, base ± rounded,
margin (unequally) serrate, apex from acute to trun-
cate, coriaceous, 7-10 pairs of nerves, patently
hairy above, densely woolly and with longer
straight hairs on the nerves below. Inflorescences
rather lax, up to 30 cm long, with up to 12(-20)
much-branched laterals under the terminal flower,
the whole inflorescence with up to 150 or more
flowers. Bracts linear or 3-partite, hairy. Pedicels
up to l(-2) cm long, woolly, with bristles and
curved prickles. Hypanthium 4-5 mm across,
densely hairy and with bristles outside. Sepals
ovate-triangular, 5-7.5 by 3-4.5 mm, entire,
shortly acuminate, woolly and with longer hairs
and at base also with bristles outside. Petals ob-
ovate to spathulate, up to 10 by 5 mm, early fal-
ling, rounded or acute, hairy, white. Stamens 30-
40, filaments up to 2 mm, anthers c. 0.5 mm
long. Pistils 100-150, ovaries on the back with
many long straight hairs, especially near apex, on
an elevated, densely hairy torus, style up to 2 mm
long, hairy at base. Collective fruits ovoid to glo-
bose, up to 8 by 10 mm, sepals ultimately slight-
ly spreading. Fruits up to 1.5 by 0.8 mm (dry),
exocarp hairy, yellow to orange, mesocarp only a
thin layer when dry.
Distribution - Continental Asia from India to
China and Vietnam, Sri Lanka; Malesia: Philip-
pines (Luzon). Introduced and naturalized in Java,
Hawaii, Jamaica, Puerto Rico, Africa, Australia,
and maybe elsewhere.
Habitat - In Luzon a species of oak and pine for-
est and in secondary growth, alt. 1000-2400 m.
Note - Root nodules from plants collected in
Java showed nitrogenase activity and can be sup-
posed to fix nitrogen under normal conditions. See
J.H. Becking, Plant and Soil 53 (1979) 541-545.
14. Rubus ferdinandi-muelleri Focke, Abh.
Naturw. Ver. Bremen 13 (1895) 165; P. van
Royen, Phan. Mon. 2 (1969) 21, f. 2, pi. 1;
Alpine Fl. New Guinea 4 (1983) 2464. —
Rubus ferdinandi Focke, Bibl. Bot. 72 (1911)
162, nom. superfl. — Type: MacGregor s.n.,
Papua New Guinea, not seen.
Rubus laeteviridis P. van Royen, Phan. Mon. 2
(1969) 29, f. 5. — Type: Womersley & van
Royen NGF 5901 = van Royen 4332, Wau.
Rubus woitapensis P. van Royen, Phan. Mon. 2
(1969) 39, f. 7. — Type: van Royen NGF
20287, Woitape-Kosibi.
Usually erect shrublets, sometimes climbing or
scrambling. Stems up to 1.5 m, densely hairy to
glabrous, prickles usually rather many, slender and
spine-like, up to 1.5 cm long, reddish to purple.
Glands sessile, red or yellow, sometimes present
on many parts of the plant. Leaves imparipinnate,
with 3-8(-9), usually opposite pairs of leaflets,
up to 13 cm long, petiole 1-3.5 cm long. Stipules
linear to lanceolate, 3-10 by 0.2-3 mm, entire
Kalkman
Rosaceae
261
or with small teeth, acute to acuminate, glabrous.
Leaflets ovate to elliptic, 1-5 by 0.5-2 cm, ter-
minal one larger than lateral ones, base rounded to
cuneate, margin (bi)serrate, apex acute, papyrace-
ous to pergamentaceous, 5-9 pairs of nerves, up-
per surface glabrous or with short appressed hairs
between and parallel with the lateral nerves, lower
surface glabrous or soft-hairy on main nerves,
sometimes with spines on the midrib. Inflores-
cence loosely branched with one or two 3- to 1-
flowered cymes under the terminal flower, up to 5
cm long. Bracts stipule-like or 3-partite. Pedicels
up to 3 cm long, hairy to glabrous, with spines.
Flowers usually erect- Hypanthium c. 4 mm across,
sparsely hairy to glabrous outside and with 1-5
purple spines alternating with the sepals. Sepals
narrowly triangular, 5-9 by 1.5-3 mm, entire,
(gradually) acuminate, indumentum outside as
hypanthium and woolly on the covered parts.
Petals obovate or elliptic to suborbicular, up to
9(-10) by 7 mm, early falling, rounded, glabrous
or with few hairs, white. Stamens 30-35, fila-
ments up to 3 mm, anthers c. 0.5 mm long. Pis-
tils 120-180, ovaries glabrous or dorsally with
some hairs, on an elevated, glabrous torus, style
up to 2 mm long. Collective fruits ovoid to sub-
globose, up to 1 cm, sepals ultimately spreading.
Fruits 1.5 by 1 mm, bright to dark red, mesocarp
not very juicy and only a thin layer when dry.
Distribution - New Guinea, New Britain.
Habitat - Clearings in forest, forest edges,
stream-banks, along tracks and roads, alt. (1550-)
1 800- 3000(- 3465) m.
Note - The complex formed by R. montis-wil-
helmi, R. papuanus, and R. ferdinandi-muelleri
must be studied with biosystematical methods. The
present delimitation of these three species is admit-
tedly provisional.
15. Rubus fraxinifolius Poiret in Lam., En-
cycl. Meth. 6 (1806) 242; Blume, Bijdr. (1826)
1107; Miq., Fl. Ind. BaL I, 1 (1855) 376; Elmer,
Leafl. Philipp. Bot. 2 (1908) 460; Merr., Enum.
Philipp. Flow. PI. 2 (1923) 227; Backer &
Bakh. f., Fl. Java 1 (1964) 514; P. van Royen,
Phan. Mon. 2 (1969) 45; Steenis, Mount. Fl.
Java (1972) pi. 45-3. — Type: Commerson s.n.,
Java.
[Rubus moluccus parvifolius Rumph., Herb. Am-
boin. 5 (1747)88, t. 47, 1.]
Rubus celebicus Blume, Bijdr. (1826) 1 107. — Ru-
bus fraxinifolius Poir. subsp. celebicus (Blume)
Focke, Bibl. Bot. 72 (1911) 151. — Type: (?)
Reinwardl s.n. in L.
Rubus fraxinifolius Poir. var. haightii Elmer, Leafl.
Philipp. Bot. 2 (1908) 461. — Type: (?) M earns
BS 4459, Pauai.
Rubus merrillii Focke, Bibl. Bot. 72 (1911) 153;
Merr., Enum. Philipp. Flow. PI. 2 (1923) 228.
— Type: Merrill BS 862 or BS 6637, Pauai.
Erect, rarely semi-scandent shrubs, up to 3 m
high. Stems glabrous, unarmed or prickles few,
straight, up to 6 mm. Glands (sub)sessile, some-
imes present on leaves and other parts up to the
flowers. Leaves imparipinnate, up to 27 cm long,
with up to 4 (or 5) opposite pairs of leaflets, pet-
iole 2-6 cm long. Stipules linear, 5-13 by 0.5-1
(-2) mm, entire or sparsely toothed, glabrate. Leaf-
lets elliptic to oblong or ovatish, 2-9(-12) by 1-4
(-6) cm, base usually rounded or cordate, margin
serrate, apex acute or acuminate to long-pointed,
papyraceous to pergamentaceous, (7— )10— 15(— 19)
pairs of nerves, both sides sparsely hairy mainly
on the nerves. Inflorescence lax and wide, up to
c. 20 cm long and wide, with up to 7 branches
under the terminal flower, the branches thyrsoid to
cymes, the entire inflorescence with up to 60 flow-
ers. Bracts lanceolate to 3-partite, up to c. 1 cm.
Pedicels up to 5 cm long, glabrous, sometimes
with small prickles. Hypanthium 5-6 mm across,
glabrous and unarmed outside. Sepals triangular,
7-13 by 3-6 mm, including the 2-5 mm long
acumen, entire, glabrous outside but woolly on
covered margins. Petals orbicular to elliptic or ob-
ovate, 7-12 by 5-9 mm, falling early, glabrous,
(greenish) white. Stamens up to more than 100,
filaments up to 3 mm, anthers c. 1 mm long.
Pistils up to more than 300, ovaries glabrous,
torus elevated, basal part without pistils and long-
hairy, upper part glabrous, style up to 1.5 mm
long. Collective fruits ellipsoid to ovoid, up to
2.5 by 1.5 cm, sepals ultimately recurved. Fruits
c. 1.5 by 0.8 mm (dry), (orange-)red, mesocarp a
thin layer when dry.
Distribution - Taiwan; Malesia: Borneo, Java,
Philippines, Celebes, Lesser Sunda Islands, Moluc-
cas, New Guinea; Solomon Islands, Bismarck Ar-
chipelago.
Habitat - Forest borders and open places in for-
est, riverbanks, deserted gardens, roadsides, and other
more or less open places, altitude 0-2500 m.
Uses - Although many collectors report on
their labels that the fruits are tasteless or worse,
they are at least in Java collected from the wild and
sold for consumption. Heyne, Null. PI. Indon.
(1950) 693, mentions the use of the leaves in cases
of slimy faeces (? dysentery, compare Rubus mo-
luccanus).
262
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Note - Erinea, as described under R. rosifolius
(p. 266), also occur in the present species.
16. Rubus lineatus Reinw. ex Blume, Bijdr.
(1826) 1108; Miq., Fl. Ind. Bat. I, 1 (1855)
378; Focke, Bibl. Bot. 72 (1910) 47, incl. var.
diengensis Focke; Backer & Bakh. f., Fl. Java 1
(1964) 514; Steenis, Mount. Fl. Java (1972)
pi. 45-4; Kalkman, Blumea 29 (1984) 322. —
Type: Reinwardt s.n., Java.
Rubus pulcherrimus Hook., Ic. Plant 8 (1845) 729,
730. — Type: Lobb s.n., Java.
Rubus lineatus Reinw. ex Blume forma pulcher-
rimus Focke in Hallier, Meded. Rijksherb. 14
(1912) 39. — Types: Elbert 1087, 1681, Lom-
bok, Griindler 2342, Sumbawa.
Shrubs, up to c. 3 m high, sometimes ± climb-
ing and up to 10 m. Stems densely long-hairy,
prickles very few or none, up to 3 mm. Leaves
pedately 5-foliolate, in the inflorescence some-
times fewer leaflets, young leaflets folded length-
wise, petiole 2-10 cm long. Stipules on the junc-
tion of twig and petiole, oblong to lanceolate, 2-4
by 0.5-1 cm, entire, cuspidate, hairy outside, fal-
ling early. Leaflets oblong to lanceolate, terminal
ones 7-18 by 2-7.5 cm, lateral ones smaller, base
acute, margin caudately serrate, apex acuminate to
caudate, pergamentaceous, (20-)30-40(-50) pairs
of nerves, upper surface variously hairy, lower
surface always densely sericeous on main nerves,
either short- woolly and also long-sericeous or quite
glabrous between the nerves, the indumentum sil-
very. Inflorescences terminal and lateral thyrsi, up
to 5 cm long and with up to 15 flowers. Bracts
stipule-like. Pedicels up to 2 cm, densely sericeous.
Hypanthium 5-9 mm across, densely sericeous out-
side. Sepals (ovate-)triangular, 6-13 by 2-7 mm,
entire, long-pointed to acuminate, indumentum out-
side as hypanthium. Petals obovate to ± rhomboid,
4-5 by 2-3.5 mm, early falling, rounded,
(greenish) white. Stamens 50-150, filaments up
to 4 mm, anthers 0.7-1 mm long. Pistils c. 80 to
over 100, ovaries long-hairy in apical part, on ele-
vated, hairy torus, style up to 5 mm long, long-
hairy. Collective fruits globose-ovoid, c. 1 cm in
diam., sepals upright to slightly spreading. Fruits
up to 2.5 by 2 mm (dry), exocarp hairy, orange to
red, mesocarp juicy but only a thin layer when dry.
Distribution - Himalayas (Nepal to Arunachal),
S China, Burma, Vietnam; Malesia: Sumatra, Bor-
neo, Java, Lesser Sunda Islands.
Habitat - Lighter places in different forest types
and in places like streambanks, landslides, roadsides,
and shrubland, altitude 1400-3000(-3800) m.
Notes - In this treatment R. lineatus has been
transferred from subg. Malachobatus (as in Focke,
I.e. 1910, and Kalkman, I.e.) to subg. Idaeobatus.
It is true that its stipules are not placed on the base
of the petiole, usual in the latter subgenus, but
rather on the junction of twig and petiole. However,
its inflorescence is rather out of tine in Malacho-
batus, not being a compound raceme but a dicha-
sium or thyrse with di- to monochasial laterals.
The species seems to be most closely related to
R. alpestris and R. neo-ebudicus. Its relationship to
the former is also apparent from transitional speci-
mens, probably hybrids, found in Borneo (R. line-
ato-alpestris Naruhashi & Sato, J. Phytogeogr.
Taxon. 32, 1984, 102) and in Java (see Kalkman,
l.c).
Rubus satotakashii Naruhashi & Cheksum, J.
Phytogeogr. Taxon. 32 (1984) 99, was interpreted
by its authors as a hybrid with R. lowii, which -
in view of the latter's relationship with R. alpes-
tris - is certainly not improbable. Tawan, Sato &
Naruhashi, J. Phytogeogr. Taxon. 39 (1991) 31,
saw intermediate characters in the supposed hybrid,
but also some unique ones.
17. Rubus lorentzianus Pulle, Nova Guinea 8
(1912) 647; P. van Royen, Phan. Mon. 2 (1969)
54; Alpine Fl. New Guinea 4 (1983) 2474. —
Type: von Romer 1276, Hellwig Mts.
Climbing or scrambling shrubs, up to 4 m high.
Stems long-hairy, glabrate, prickles many, straight,
stout, up to 9 mm long, red. Leaves 3-foliolate,
upper ones sometimes simple, petiole 1-5 cm
long. Stipules ovate to lanceolate, usually oblique,
10-18 by 3-15 mm, entire to serrate, acute to
caudate, hard, sometimes with prickles. Leaflets
obovate, 2-7 by 1.5-3.5 cm, lateral ones shorter
and relatively broader, margin serrate, apex rounded,
rarely acute to acuminate, very stiff coriaceous,
4-8 pairs of nerves, both sides slightly hairy on
main nerves when young, hairs often disappearing
with age, leaflets often folded along the midrib. In-
florescence with up to 4 cymes below the terminal
flower, cymes with 1-3 flowers, the whole inflo-
rescence usually with less than 8 flowers. Bracts
stipule-like. Pedicels up to 3 cm, hairy and with
some prickles. Hypanthium up to 7 mm across,
short-hairy and with many prickles outside. Sepals
ovate to triangular, 9-14 by 5-8 mm, entire,
caudate, rather sparsely hairy and with many long
prickles outside, woolly on covered margins. Petals
obovate to suborbicular, 8-1 1 by 6.5-9 mm, round-
ed, white. Stamens 30-45, filaments up to 5 mm,
anthers c. 1 mm long. Pistils 15-45, ovaries gla-
Kalkman
Rosaceae
263
brous, on elevated, hairy torus, style up to 4 mm
long. Collective fruits ovoid, up to 1.5 cm across.
Fruits up to 4.5 by 3 mm when dry, orange to red,
mesocarp fleshy.
Distribution - New Guinea.
Habitat - Forest edges, openings in forest, shrub-
land, sometimes in grassland, altitude 2200-3650
(-3890) m.
Note - High-polyploid with 2n = 126, according
to one count by Borgmann, Zs. f. Bot. 52 (1964)
144, sub Rubus spec.
18. Rubus lowii Stapf in Hook., Ic. Plant. 23
(1894) L 2289; Trans. Linn. Soc. BoL 4 (1894)
145; Naruhasi & Sato, Tukar-Menukar 2 (1983)
14, incl. var. panalabanensis Naruhashi & Sato,
nom. nud.; Naruhashi et al., J. Phytogeogr.
Taxon. 32 (1984) 102, f. 3, incl. var. panalaba-
nensis Naruhashi & Sato, descr. — Type: Low
s.n., Mt Kinabalu.
Climbing or scrambling shrubs. Stems up to
6 m, rather densely hairy, glabrate, prickles absent
or few, stout. Glands sessile or stalked, sometimes
present on all parts of the plants. Leaves 3-folio-
late, upper ones sometimes simple, petiole 0.7-2
cm long. Stipules lanceolate, 6-13 by 1.5-5 mm,
long persistent, acute, entire or with small teeth,
glabrous. Leaflets elliptic to elliptic-ovate, terminal
ones 2-7 by 1-4 cm, lateral ones slighdy smaller,
base acute, margin serrate, apex acute to acuminate,
coriaceous, 5-8 pairs of nerves, both surfaces
with long hairs on main nerves, upper surface rare-
ly also with hairs between nerves. Inflorescence
with up to 4 few-flowered, axillary cymes below
the terminal flower. Bracts stipule-like. Pedicels
up to 3 cm long, hairy. Hypanthium c. 6 mm
across, short-hairy outside. Sepals triangular to
ovate, 8-11 by 4-6 mm, after anthesis slightly
growing, ± entire, caudate, acumen up to 3 mm
long, sparsely hairy outside and with woolly cov-
ered margins. Petals obovate to suborbicular, 6-7
by 4-5.5 mm, rounded, white (to pinkish?). Sta-
mens 30-45, filaments up to 5 mm, anthers c. 1
mm long. Pistils 15-25, ovaries glabrous, on
slightly elevated, hairy torus, style up to 6 mm
long. Collective fruits ovoid, c. 1.5 by 1 cm.
Fruits up to 4 by 3 mm, red, mesocarp fleshy,
tough when dry.
Distribution - Borneo, only seen from Mt Kina-
balu.
Habitat - In open forest, forest edges, shrubland,
altitude 3000-3960 m.
Note - Closely related to, maybe even conspec-
ific with R. macgregorii from Celebes and New
Guinea. The latter species has distinctly armed
stems and hairy ovaries. Both are also related to R.
alpestris with 5-foliolate leaves.
19. Rubus macgregorii F. Muell., Trans. Roy.
Soc. Vict. 1, 2 (1889) 4; Steenis, Bull. Jard.
Bot. Buitenzorg III, 13 (1934) 245; Merr. &
Perry, J. Arnold Arbor. 21 (1940) 179; P. van
Royen, Phan. Mon. 2 (1969) 52; Alpine Fl. New
Guinea 4 (1983) 2472. — Type: McGregor s.n.,
Mt Victoria, Papua New Guinea.
Creeping or scrambling small shrubs. Stems
sparsely soft-hairy, glabrate, prickles rather few,
curved, up to 2(-4) mm long. Leaves 3-foliolate,
upper ones sometimes simple, petiole 1-5 cm
long. Stipules elliptic to elliptic-lanceolate, 5-12
by 1-5 mm, entire or toothed, acuminate to cau-
date, ± glabrous. Leaflets obovate to elliptic, ter-
minal ones 1.5-3.5 by 1-2.5 cm, lateral ones
smaller, base narrowed, margin serrate, apex round-
ed with or without a short acumen, stiff coriaceous,
4-7 pairs of nerves, both surfaces very sparsely
hairy on main nerves. Inflorescences poor, often
only the terminal flower, sometimes 1 or 2 flow-
ers below it. Bracts stipule-like. Pedicels up to
2 cm long, hairy and with some curved prickles.
Hypanthium 5-6 mm across, sparsely hairy out-
side and with some straight, short prickles. Sepals
ovate to triangular, 8-12 by 5-7 mm, growing
after anthesis, entire, acuminate to 4 mm caudate,
long-hairy outside and woolly on the covered parts.
Petals obovate to orbicular, 8-9 by 7-8.5 mm,
rounded, white. Stamens 40-50, filaments up to
4 mm, anthers c. 1 mm long. Pistils 30-40, ova-
ries long hairy on the dorsal side and at the top, on
an elevated, hairy torus, style up to 3.5 mm long,
with hairs at base. Collective fruits ovoid, up to
2 cm across. Fruits up to 4 by 3 mm, purple (?),
mesocarp fleshy, a thin layer when dry.
Distribution - Celebes, Papua New Guinea.
Habitat - Thickets in grassland, altitude 2600-
3600 m.
Note - See note under the related R. lowii.
20. Rubus montis-wilhelmi P. van Royen,
Phan. Mon. 2 (1969) 19, f. 1; Alpine Fl. New
Guinea 4 (1983) 2462. — Type: Millar & van
Royen NGF 14645, Mt Wilhelm.
Rubus keysseri Schltr. ex Diels, Bot. Jahrb. 62
(1929) 481. See note.
Erect shrublets, ± climbing or straggling when
larger, up to 1(-1.5) m high. Stems sparsely hairy,
glabrate, prickles spine-like, slender, up to 1(-1.5)
264
Flora Malesiana ser. I, Vol. 11 (2) (1993)
mm long, reddish. Glands (sub)sessile, red or yel-
low, usually scattered on many parts of the plant.
Leaves up to 12 cm long, bipinnate to pinnate (to
the apex) or sometimes tripinnate (at very base),
primary pinnae 4-10, (sub)opposite, petiole 1-3
cm long. Stipules linear-lanceolate, 5-12 by 0.5-
1.5 mm, (sub)glabrous. Leaflets 3-5 pairs on the
primary pinnae, ovate to ovate-elliptic or elliptic-
oblong, 2-10 by 1.5-6 mm, base acute, margin
serrate to pinnatipartite, apex usually acuminate,
pergamentaceous, with 3-5 pairs of nerves, both
surfaces glabrous or with few hairs. Inflorescence
loosely branched with one or two 1- to 3-flowered
cymes under the terminal flower, up to 5 cm long.
Bracts linear, up to 3 mm. Pedicels up to 2 cm
long, sparsely hairy and with spines. Flowers usu-
ally (sub)pendulous. Hypanthium up to 4.5 mm
across, sparsely hairy and with some spines out-
side, the largest ones almost as long as and alter-
nating with the sepals. Sepals narrowly triangular,
5-9 by 2-3.5 mm, long-pointed, entire, indu-
mentum outside as hypanthium and very shortly
woolly on the covered parts. Petals obovate to el-
liptic, falling early, up to 12 by 10 mm, rounded,
white. Stamens 25-35, filaments up to 3 mm,
anthers 0.5-0.8 mm long. Pistils 100-150, ova-
ries glabrous, on elevated, glabrous torus, style up
to 1.5 mm long. Collective fruits ovoid to ellip-
soid, up to 1 cm across, compact, sepals ultimately
spreading. Fruits 1.5 by 1 mm, (dark) red, meso-
carp not very juicy.
Distribution - Papua New Guinea.
Habitat - In (the edges of) subalpine and alpine
shrubland and forests, altitude 2660-3660 m.
Note - Probably easily hybridizing with R.fer-
dinandi-muelleri and R. papuanus, see there. Rubus
keysseri Schltr. was based on what looks like a
hybrid specimen with the first-mentioned species
(isotype seen from BM).
21. Rubus neo-ebudicus Guillaumin, J. Arnold
Arbor. 12 (1931) 249. — Type: Kajewski 249,
Tanna I.
Rubus brassii Merr. & Perry, J. Arnold Arbor. 21
(1940) 182; Zandee & Kalkman, Blumea 27
(1981) 105. — Type: Brass 2891, San Christo-
bal.
Straggling or climbing, unarmed shrubs, up to
4 m high. Stems shortly woolly to glabrous.
Leaves pedately 5-foliolate, the upper ones often
3-1-foliolate, petiole 2-5 cm long. Stipules
(linear-)lanceolate, 5-10 by 1-3 mm, entire, gla-
brous or with hairs. Leaflets (oblong-)lanceolate,
rarely ovate-lanceolate, terminal leaflet 8-14 by
2-4.5 cm, on 2-8(-15) mm long petiolule, lat-
eral leaflets smaller, sessile or to 2 mm petioluled,
base acute, margin mostly biserrate, apex acumi-
nate, herbaceous, (16-) 18-21 (-26) pairs of nerves,
sparsely hairy on both surfaces. Inflorescence lax,
with up to 5 dichasial, up to 8-flowered branches
under the terminal flower, up to 12 cm long.
Bracts up to 1 cm, often toothed, those under the
lateral dichasia usually 3-partite. Pedicels up to 2
cm long, all axes shortly woolly to glabrous.
Hypanthium 5-6 mm across, sparsely hairy and
sometimes with few short glandular hairs outside.
Sepals narrowly triangular, 8-13 by 3-5 mm,
acuminate to caudate (acumen up to 4 mm), entire,
few hairs outside and covered margins woolly.
Petals obovate to elliptic, 9-12 by 6-9 mm, fal-
ling early, obtuse, white. Stamens 45-100, fila-
ments up to 4 mm, anthers c. 1 mm long. Pistils
80-100 or more, ovaries glabrous or with few
hairs, on elevated, sparsely hairy torus, style up to
2.5 mm long. Collective fruits ovoid, up to 1 cm
across, sepals upright. Fruits 2 by 1 mm when dry,
orange to red, mesocarp only a thin layer when dry.
Distribution - New Ireland, New Britain, Solo-
mon Islands, New Hebrides.
Habitat - Forest, altitude (180-)600-1700 m.
Uses - According to Woodley (ed.), Medicinal
Plants of Papua New Guinea I. Morobe Prov. (1991)
120 (sub R. brassii) the extracted stem sap is drunk
as a tonic by elderly people. Other species are, ac-
cording to this source, used for the same purpose
elsewhere.
Notes - Mr. P.S. Green (Kew) drew my atten-
tion to the conspecificity of the species reported
from the New Hebrides with R. brassii from New
Britain and the Solomon Islands.
The relationships of the present species seem to
lie with R. lineatus and R. alpestris, which is pecu-
liar since both species are absent from New Guinea
and the Pacific.
22. Rubus niveus Thunb., Diss. Rubo (1813)
9, f. 3; Merr., Enum. Philipp. Flow. PI. 2
(1923) 229; Backer & Bakh. f., Fl. Java 1 (1964)
515; Lauener & Ferg., Not. Roy. Bot. Gard.
Edinb. 30 (1970) 276; Steenis, Mount. Fl. Java
(1972) pi. 45-5. — Non Rubus niveus Wall, ex
Hook, f., Fl. Brit. India 2 (1878) 335 = Rubus
hypar gyrus Edgew. — Type: Thunberg s.n.,
Java.
Rubus horsfieldii Miq., Fl. Ind. Bat. I, 1 (1855)
375, t. 7; Koord., Nat. Tijd. Ned. Indie 60 (1901)
276. — Rubus niveus Thunb. subsp. horsfieldii
(Miq.) Focke, Bibl. Bot. 72 (1911) 183. — Type:
Horsfield s.n., Java.
Kalkman — Rosaceae
265
Usually erect, sometimes climbing shrubs, up to
2 m high, the often drooping branches up to 3.5 m
long. Stems sparsely hairy, glabrescent, prickles
usually rather few, straight to curved, up to 7 mm.
Leaves imparipinnate, up to 27 cm long, petiole
1.5-5 cm long. Stipules (linear-)lanceolate, 6-16
by 1— 3(— 5) mm, slightly hairy to glabrous. Leaf-
lets 2-4(-5) opposite pairs, elliptic or rhombic
to ovate, sometimes ovate-lanceolate, 2-8 by 1-4
cm, base usually acute, margin serrate to biserrate
but the basal part often entire, apex acute to acu-
minate, papyraceous to pergamentaceous, 6-9
pairs of nerves, soft-hairy above but soon glabrate,
lower surface with a woolly silvery-white felt of
short curly hairs all over and with longer straight
hairs on main nerves. Inflorescence usually rich
and branched, a compound leafy thyrse up to 20
cm long. Bracts stipule-like. Pedicels up to 1.5
cm, hairy. Hypanthium 2-3 mm across, hairy
outside. Sepals triangular, 4-7 by 1.5-2.5 mm
including the up to 2 mm long acumen, densely
woolly outside, acumen mostly glabrous. Petals
falling rather early, suborbicular, 3.5-5 by 3-3.5
mm, 1 mm clawed, pink. Stamens 25-35, fila-
ments up to 4 mm, anthers c. 0.5 mm long. Pis-
tils 50-75 or more, ovaries rather densely long-
hairy, on elevated, hairy torus with pistils down to
the base, style up to 3 mm long. Collective fruits
globular to broadly ovoid, up to c. 1 cm across,
compact, sepals spreading. Fruits c. 2.5 mm long,
exocarp densely hairy, red but the colour masked
by the dark (blue to blackish) hair-cover, mesocarp
only a thin layer when dry.
Distribution - Continental Asia from Kashmir
to Vietnam, Sri Lanka, Taiwan; Malesia: Sumatra,
Java, Lesser Sunda Islands, Luzon, Celebes. Intro-
duced and naturalized in Southern and Eastern Africa
(Stirton, Bothalia 13, 1981, 346). Introduced and
cultivated in peninsular Malaysia where it became
naturalized on Fraser's Hill, Pahang. Introduced
and (recently) cultivated near Kainantu, Eastern
Highlands Prov., Papua New Guinea (R.H. Con-
verse, in litt. 1986). Also cultivated for the fruits
in Florida, USA ('Mysore raspberry'), maybe also
elsewhere.
Habitat - Open and half-shaded places like
hedges, shrubland, grassfields, abandoned gardens,
roadsides, Eucalyptus savannas, rarely in forest,
altitude (600-) 1000- 2900 m.
Uses - Fruits edible, see also under Distribu-
tion.
Notes - The species is on the Asian continent
more variable than in Malesia where infraspecific
taxa cannot be recognized. The synonymy given
above is incomplete for the continent.
In some of the specimens from Timor the wool-
ly indumentum on the underside of the leaflets is
missing, but there are dots and patches of dense
long hairs, possibly galls.
23. Rubus papuanus Schltr. ex Diels, Bol Jahrb.
62 (1929) 481; Merr. & Perry, J. Arnold Arbor.
21 (1940) 182; P. van Royen, Phan. Mon. 2
(1969) 26, f. 4; Alpine Fl. New Guinea 4 (1983)
2470. — Type: Keysser 36, lost; neotype: Brass
4246, Mt Albert Edward.
Erect or scrambling small shrubs, up to 80
cm high. Stems soft-hairy, glabrescent, prickles
straight, slender, spine-like, up to 1 cm, red. (Sub)-
sessile red or yellow glands often present on many
parts. Leaves imparipinnate, up to 10 cm long,
petiole 0.5-1 (-1.5) cm long. Stipules oblong to
linear- lanceolate, 4-9 by 1-3 mm, glabrous. Leaf-
lets 6-9(-10) (sub)opposite pairs, (broadly) ovate,
6-15 by 4-9 mm, base rounded to cuneate, mar-
gin rather deeply serrate, apex pointed to acumi-
nate, pergamentaceous, 3-5 pairs of nerves, both
surfaces glabrous or almost so. Inflorescence poor,
at most 1 or 2 axillary flowers under the terminal
one. Pedicels up to 2.5 cm long, soft-hairy. Flow-
ers more or less pendulous. Hypanthium up to 6
mm across, soft-hairy to glabrous outside and with
some spines, the largest of those up to c. 1 cm,
alternating with the sepals. Sepals narrowly trian-
gular, 6-1 1 by 2-4 mm including the up to 3 mm
long acumen, indumentum outside as hypanthium
and woolly on the covered parts. Petals falling ear-
ly, broadly obovate to orbicular, 6— 12(— 18) mm
long, rounded or retuse, white. Stamens 20-50,
filaments up to 3 mm, anthers c. 0.8 mm long.
Pistils 70-100, ovaries glabrous or dorsally with
few hairs, on elevated, glabrous torus with pistils
down to the base, style up to 1.5 mm long. Col-
lective fruits ovoid, up to 2 by 1.5 cm, sepals ul-
timately spreading. Fruits up to 2.5 mm long,
bright red, mesocarp fleshy but only a thin layer
when dry.
Distribution - New Guinea, New Ireland. In
1965 introduced in England (Grasmere, Westmor-
land) and there it is hardy.
Habitat - Subalpine and alpine shrubland, grass-
lands, open places in forest, and forest edges, alti-
tude (2100-)3000-3650 m.
Note - This may be a high altitude form of R.
ferdinandi-muelleri. See the notes there and to R.
montis-wilhelmi.
24. Rubus rosifolius J.E. Smith, PI. Icon.
Hact. Ined. 3 (1791), t. 60, 'rosaefolius' as in
266
Flora Malesiana ser. I, Vol. 11 (2) (1993)
many other references; Blume, Bijdr. (1826)
1107; Hook., Ic. Plant. 4 (1840) t. 349; Miq.,
Fl. Ind. Bat. I, 1 (1855) 375; Elmer, Leafl.
Philipp. Bot. 2 (1908) 462; Merr., Enum. Phi-
lipp. Flow. PI. 2 (1923) 230; Ochse, Fruits
(1931) 107, pi. 42; Backer & Bakh. f., Fl. Java 1
(1964) 515, excl. subsp. sumatranus Focke; P.
van Royen, Phan. Mon. 2 (1969) 34; Steenis,
ML Fl. Java (1972) pi. 45-3; Kalkman in Steenis
(ed.), Blumea 28 (1982) 168 (reduction of Gilli
names). — Type: Commerson s. n., Mauritius.
Rubus rosifolius J.E. Smith var. coronarius Sims,
Curt. Bot. Mag. (1816) t. 1783; Backer &
Bakh. f., Fl. Java 1 (1964) 515. — Type: plate
and description in Sims, I.e.
Rubus javanicus Blume, Bijdr. (1826) 1108. —
Type: Blume 1571, Java.
Rubus tagallus Cham. & Schlechtend., Linnaea 2
(1827) 9; Elmer, Leafl. Philipp. Bot. 2 (1908)
461; Merr., Enum. Philipp. Flow. PI. 2 (1923)
230. — Type: de Chamisso s.n., Luzon.
Rubus apoensis Elmer, Leafl. Philipp. Bot. 5
(1913) 1618. — Type: Elmer 10464, Mindanao.
Rubus mingendensis Gilli, Ann. Naturhist. Mus.
Wien 83 (1980) 457, incl. var. trichocarpa Gilli,
nom. nud., inval. — Type: Gilli 111, Papua
New Guinea, Chimbu.
Rubus x dosedlae Gilli, I.e. 456. — Type: Dosedla
45 A, Papua New Guinea, Mt Hagen.
Erect or scrambling, rarely climbing shrubs, up
to l(-3) m high. Stems soft-hairy, glabrescent,
prickles usually rather few, curved to straight, 1-5
mm. Sessile, pale yellow glands usually present
on many parts of the plants. Leaves imparipinnate,
up to 18 cm long, petiole 1-5.5 cm long. Stipules
linear, 4-9 by 0.5-1 mm, entire. Leaflets in (1-)
2_3(-4) opposite pairs, ovate to ovate-oblong,
sometimes elliptic to oblong, 2-6 by 1-2.5 cm,
terminal leaflets up to 8 by 4 cm, base acute to
cordate, margin biserrate, apex acute to long-taper-
ing, papyraceous, (4-)7-9(-ll) pairs of nerves,
both surfaces soft-hairy. Inflorescence with up to
4 dichasia in the axils of the upper (reduced) leaves,
rarely longer than 10 cm, with up to 10 flowers.
Pedicels up to 4 cm long, hairy. Hypanthium 4-6.5
mm across, with scattered hairs and many glands
outside. Sepals ovate to narrowly triangular, 7-15
(-22) by 2-5 mm, including the up to 5 mm
acumen, entire, indumentum outside as hypanthium
and shortly woolly on the covered margins. Petals
falling early, broadly obovate to ovate, 8-17 by
6-12 mm, obtuse, white. Stamens 60-140, fila-
ments up to 8 mm, anthers 0.5-0.8 mm long.
Pistils up to c. 600, ovaries with some apical hairs
and usually many shortly stalked pale glands, on
elevated, hairy torus with pistils down to the base,
style up to 2 mm long. Collective fruits ovoid to
globose or ellipsoid, up to 2.5 cm across, sepals
recurved. Fruits c. 1.5 mm long when dry, red,
mesocarp juicy, only a thin layer when dry.
Distribution - Continental Asia (Assam, Cam-
bodia, Vietnam), Japan (?), Taiwan, New Britain,
New Ireland, New Hebrides, New Caledonia, Aus-
tralia (Queensland, New South Wales); in Malesia:
Borneo, Java, Philippines, Celebes, Lesser Sunda
Islands, New Guinea, Bougainville. Introduced and
naturalized in many parts of the world: Africa, C &
S America, island groups in the Pacific and Indian
Ocean.
Habitat - In open (secondary, anthropogenic)
places like clearings, forest-edges, roadsides, land-
slides, grassland, riverbanks, fallow gardens, also
in shrubland and rarely in the undergrowth of lighter
types of forest, altitude 0-2000(-2400) m, in
Celebes also reported from 2800-2900 m.
Uses - Because of its easy growth, nice flowers
and edible fruits often cultivated in sunny gardens,
within and outside its natural area. Quisumbing,
Medic. PI. Philipp. (1951) 354, records the use of
a decoction of roots as an expectorant.
Notes - Garden-forms exist with more than 5
petals in the flower, which resembles a small rose.
They are usually called var. coronarius Sims. These
plants may produce fruits and have sometimes es-
caped from cultivation.
Some specimens possess remarkable erinea, con-
sisting of dots of a very dense indumentum on the
leaves, probably caused by the gall-mite Eriophyes
rubierineus. See Docters van Leeuwen, Zoocecidia
(1926) 220. The same kind of erinea is also found
in some other species.
25. Rubus sumatranus Miq., Sumatra (1861)
307; Lauener & Ferg., Not. Roy. Bot. Gard.
Edinb. 30 (1970) 280. — Rubus rosifolius I.E.
Smith subsp. sumatranus (Miq.) Focke, Bibl.
Bot. 72 (1911) 155; Backer & Bakh. f., Fl. Java
1 (1964) 515. — Type: Teijsmann s.n., Sumatra.
Rubus asper auct. non Wall, ex D. Don: Focke,
Bibl. Bot. 72 (1911) 157, f. 67; Backer, Schoolfl.
Java (191 1)454.
Erect or scrambling to semi-scandent shrubs, up
to 2 m high. Stems with many up to 5 mm long
gland-tipped, reddish, setose hairs and usually also
with soft, curly hairs, prickles usually not many,
curved, up to 5 mm. Sessile, pale or red globular
glands often present on many parts of the plant,
especially the leaves. Leaves imparipinnate, up to
Kalkman — Rosaceae
267
1 mm
Fig. 3. Rubus sumatranus Miq. a. Branch with old flowers; b. stem with prickles and glandular hairs;
c. collective fruit (a, b: Lorzing 4753; c: Kochummen FRI 16456).
21 cm long, petiole (1— )2 — 6 cm long. Stipules
linear, 3-6 by up to 0.5 mm, entire. Leaflets in
2-3(-4) opposite pairs, oblong to oblong-ovate,
2.5-7 by 0.8-2 cm, base acute to rounded, mar-
gin serrate to biserrate, apex acute to long-taper-
ing, herbaceous, 7-12 pairs of nerves, soft-hairy
on both surfaces and with gland-tipped long hairs.
Inflorescence consisting of up to 5(-8) cymes in
the axils of the upper leaves, up to 25 cm long,
with up to 20 flowers. Bracts in the cymes often
leaf-like. Pedicels up to 4 cm long. Hypanthium
4-5 mm across, with some soft hairs and with
gland-tipped setose hairs outside. Sepals narrowly
triangular, 7-14 by 2-3.5 mm, including the up
268
Flora Malesiana ser. I, Vol. 11 (2) (1993)
to 5 mm long acumen, indumentum outside as
hypanthium and also woolly on the covered mar-
gins. Petals falling early, oblong to obovate, 8-10
by 2-4 mm, obtuse, slightly fimbriate at apex,
white. Stamens up to c. 120, filaments up to 4 mm,
anthers 0.5 mm long. Pistils up to c. 500, ovaries
glabrous, on elevated, glabrous torus with pistils
down to the base, style up to 2 mm long. Collec-
tive fruits ellipsoid, up to 1.5 by 0.8 cm when
dry, sepals recurved. Fruits 1-1.5 mm long, orange-
red to red, mesocarp only a very thin layer when
dry. - Fig. 3.
Distribution - NE India, Thailand, Laos, Viet-
nam, S China, Taiwan, Japan; in Malesia: Suma-
tra, Peninsular Malaysia, Java.
Habitat - Clearings, roadsides, thickets, tea plan-
tations, forest borders, and similar open places, very
rarely reported from lighter types of forest, altitude
(Sumatra and Malaya) 500-2000 m.
Uses - Fruits edible, pleasant of taste.
Note - Confused with R. croceacanthus Leveille
(/?. asper Wall, ex D.Don 1825, nom. illeg., non
Presl 1822) from continental Asia.
Subgenus Malachobatus
Rubus subg. Malachobatus (Focke) Focke, Bibl. Bot. 72 (1910) 41; Kalkman, Blumea
29 (1984) 319. — Rubus sect. Malachobatus Focke, Abh. Naturw. Ver. Bremen 4
(1874) 187, 201.
Leaves mostly simple (in Malesia always), entire or lobed, usually pedately nerved
with on either side at the very base of the midrib 2 or 3 main side nerves, each with 2-5
basiscopic lateral nerves, above the base pinninerved, nerves usually terminating in the
margin, nervation more rarely palmate with 3-7 main nerves, or pinnate. Stipules free, on
the twig near the petiole-base, usually rather persistent. Inflorescences terminal, compound
racemes or thyrses, side branches racemose or di- or monochasial, the lower branches
axillary to leaves. Flowers usually bisexual, some species (gyno)dioecious. Hypanthium
saucer- to cup-shaped. Sepals subequal or inner ones distinctly narrower, entire or (usual-
ly) with up to 5 teeth on the not-covered margins. Petals in some species wanting or only
one left. Fruits cohering, falling as a whole together with the dried torus.
Distribution — Many species (c. 80?), centred in Continental Asia and Malesia, ex-
tending to Japan, Australia, and the W and SW part of the Pacific Ocean. In Malesia 19
species and some incompletely known ones (see Kalkman, I.e.). New Guinea is poor in
species belonging to this subgenus, which contrasts with the two other subgenera.
26. Rubus alceifolius Poiret in Lam., Encycl.
Meth. 6 (1806) 247, 'alcaefolius, alceaefolius',
the latter orthography also in many other refer-
ences; Blume, Bijdr. (1826) 1109; Miq., Fl. Ind.
Bat. I, 1 (1855) 379; Suppl. 1 (1860/61) 116,
308; Kuntze, Meth. Speciesbeschr. (1879) 56;
Ridley, Fl. Mai. Penins. 1 (1922) 677; Backer
& Bakh.f., Fl. Java 1 (1964) 516. — Rubus
moluccanus L. var. alceifolius (Poiret) Kuntze,
Rev. Gen. PI. 1 (1891) 222. — Type: Com-
merson s.n., Java.
Shrubs with arching or climbing branches up to
5 m long. Stems rather densely covered with patent
and straight hairs, mixed with shorter and thinner
hairs and with stalked glands, prickles usually
rather many, stout. Leaves orbicular to broadly
ovate in outline, (10-)12-26 by (9-)12-26 cm,
5-7-lobed with up to 4 cm deep incisions, lobes
rounded and shallowly lobed, base deeply cordate,
margins grossly and evenly serrate, apex obtuse to
acute, herbaceous to slightly coriaceous, nervation
pedate with 5-7 pairs of nerves, venation reticu-
late, sometimes the upper surface distinctly bullate
between the veins, upper surface hairy, lower sur-
face with a usually closed felt of short, curly hairs
and with many long, patent hairs on the nerves.
Petiole 3-11 cm long. Stipules often rather per-
sistent, orbicular in outline, up to c. 2 cm, deeply
digitately divided with the lobes pinnate, lobes
Kalkman — Rosaceae
Fig. 4. Rubus alceifolius PoireL Inflorescence and leaf. Mt Salak, Java Photo J.H. Becking.
270
Flora Malesiana ser. I, Vol. 11 (2) (1993)
thread-like, at most 0.3 mm wide, hairy outside
and on margins. Inflorescence a terminal com-
pound raceme with 12 or more laterals, the lower
of them in the axils of leaves, up to 50 cm long,
the lateral racemes up to 12 cm long, all axes ter-
minating in a flower. Bracts rather persistent, pin-
natifid to -partite, with thin lobes. Pedicels 1-1.5
cm long, densely hairy as are all axes in the inflo-
rescence. Flowers bisexual, flower buds ± globu-
lar. Hypanthium cup-shaped, 6-9 mm across,
densely woolly and with long patent hairs outside.
Sepals ovate, 6-10 by 4-7 mm, acute to acumi-
nate, not-covered margins with 2-5 teeth up to
2(_4) mm long, indumentum outside as hypan-
thium. Petals early falling, orbicular, 5.5-9.5 by
4.5-9 mm, distinctly clawed, rounded or notched
at apex, white. Stamens 160-230, filaments up
to 5 mm, anthers 0.5-0.8 mm long, with long
hairs. Pistils up to 150, ovaries glabrous, on ele-
vated, glabrous torus, style up to 10 mm long.
Collective fruits globular, c. 1 cm, sepals under
ripe fruits spreading. Fruits 2-4 by 2-3 mm
when dry, red, mesocarp juicy, a thin layer when
dry. - Fig. 4.
Distribution - China, Taiwan, Burma, Thailand,
Laos, Cambodia, Vietnam; Malesia: Sumatra, Ma-
laya, Borneo, Java, Celebes, Lesser Sunda Islands.
Introduced in Australia (Queensland), Madagascar,
and Mascarenes.
Habitat - In light places like forest edges, road-
sides, secondary forest, thickets, and riverbanks, alti-
tude (0-)500-1400(-1600) m.
Uses - Shoots are eaten (Sumatra), a kind of
use which is rarely mentioned for species of
Rubus. Roots of the species are boiled and taken
against dysentery (Malaya).
Note -Rubus alceifolius differs from all varie-
ties and forms of the related R. moluccanus in the
shape of the closed flowerbuds (globular, not ovoid-
pointed) and in the stipules which have very thin,
filiform lobes.
27. Rubus beccarii Focke, Bibl. Bot. 72 (1910)
62. — Type: Beccari 175, Sumatra.
Probably large, climbing shrubs. Stems densely
hairy, prickles rather many, rather small. Leaves
ovate, 9-11.5 by 6.5-7.5 cm, not lobed, base
shallowly cordate, margin serrate, apex acute to
shortly acuminate, rather thick and firm, nervation
pinnate or pedate, with 7 or 8 pairs of nerves, those
not always terminating in the margin, venation
transverse, upper surface long-hairy, lower surface
with many long, straight, patent hairs and with
few short, curly hairs. Petiole c. 2 cm long, hairy.
Stipules c. 1 cm long, deeply divided, lobes thin.
Inflorescence only fragmentarily known, axes den-
sely hairy. Flowers possibly functionally unisexual,
only males seen, flower buds globular. Hypanthium
c. 5 mm across, densely hairy outside with woolly
felt and long, straight hairs. Sepals broadly ovate,
up to 6 by 6 mm, uncovered margins with c. 5
small teeth, covered margins entire, indumentum
outside as hypanthium. Petals 7 by 6 mm (not
full-grown), rounded apex. Stamens c. 115, gla-
brous, anthers 1 mm long. Pistils many, not de-
veloped in male flowers, pistillodes long-hairy on
back and in basal part of style. Collective fruits
not seen.
Distribution - Only known from the type spec-
imen from Mt Singgalang in West Sumatra. Close-
ly allied to, and maybe conspecific with R. smithii.
28. Rubus benguetensis Elmer, Leafl. Philipp.
Bot. 1 (1908) 296; Merr., Enum. Philipp. Row.
PI. 2 (1923) 227. — Type: Elmer 8383, Luzon.
Climbing or scrambling shrubs. Stems up to
15 m long, young twigs (rather) densely covered
with patent and with curly hairs mixed with glan-
dular hairs, prickles few to rather many, small.
Leaves ovate to elliptic, (6-)7.5-13 by (3.5-)5-8
cm, not lobed, in sterile shoots up to 17 by 9 cm
and shallowly 3-lobed, base cordate to subtruncate,
margin evenly serrate, apex acute to shortly acumi-
nate, herbaceous to stiffly coriaceous, nervation
pinnate with up to 6 pairs of nerves, sometimes ±
pedate, venation transverse, both surfaces more or
less densely hairy at least on nerves and veins.
Petiole 0.5-l(-1.5) cm long. Stipules often rather
persistent, pinnatisect to -partite with 2-4 pairs
of lobes, up to 9 by 3 mm. Inflorescence laxly
paniculate, a compound raceme with the ultimate
branching sometimes cymose, up to c. 20 laterals
with up to 30 flowers, the lowermost laterals in
the axils of leaves, the entire inflorescence up to
c. 35 cm long, the laterals up to 15 cm. Bracts usu-
ally tripartite, up to c. 8 mm long. Pedicels 0.7-
1.5 cm long, densely hairy as are the branches of
the inflorescence. Flowers bisexual, flower buds
ovoid. Hypanthium saucer-shaped, 3.5-4 mm
across, shortly woolly outside and also with long
hairs. Sepals ovate, sharply pointed, 6-7 by 2.5-5
mm, inner ones narrower than outer ones, covered
margins entire, outer margins with usually one
minute tooth under the apex, indumentum outside
as hypanthium, the outside pink to purple as are
the pedicels. Petals none, rarely one or a semi-
petaloid stamen. Stamens 50-80, glabrous, fila-
ments up to 5 mm, anthers 0.5-0.8 mm long.
Kalkman — Rosaceae
271
Pistils (13-)20-30, glabrous, ovaries on slightly
elevated, hairy torus, style up to 5 mm long. Col-
lective fruits globular, up to 8 mm diam., sepals
closing after anthesis. Fruits 3.5-6 by 2-2.5 mm
when dry, black, mesocarp probably fleshy and
rather thick but when dry only a thin, tough layer.
Distribution - Borneo (seen from Sarawak,
Sabah), Luzon, C and S Celebes.
Habitat - Primary forest on slopes, also on open
cliffs, altitude (150-)600-2900 m.
Ecology - Several collections seen from lime-
stone but also on other soil types.
29. Rubus chrysophyllus Reinw. ex Miq., Fl.
Ind. Bat. 1, 1 (1855) 380; Kuntze, Meth. Species-
beschr. (1879) 56, 76; Backer & Bakh.f., Fl.
Java 1 (1964) 516; Steenis, Mount. Fl. Java
(1972) pi. 45-2. — Rubus moluccanus L. var.
chrysophyllus (Reinw. ex Miq.) Kuntze, Rev.
Gen. PI. 1 (1891) 222. — Types: Reinwardt
s.n., holo; Junghuhn s.n.; both Java.
Rubus moluccanus L. var. ochrascens Blume, Bijdr.
(1826) 1109. — Type: Blume s.n., Java.
Shrubs, up to 4 m high, overhanging branches
up to 10 m long. Stems with a dense yellowish
indumentum of short, curly hairs and long, (semi-)
appressed, straight or wavy hairs, glabrate, prickles
few, short, or twigs unarmed. Leaves ovate to
broadly ovate, 7-22 by 7-18 cm, shallowly 3-7-
lobed, base truncate to shallowly cordate, margins
grossly and unevenly serrate, apex acute, stiff-
coriaceous, nervation pedate with 6-9 pairs of
nerves, venation reticulate, on upper surface the
squarish intervenial fields distinctly bullately raised,
upper surface soon glabrous, lower surface with a
dense, closed felt of short, curly hairs and on nerves
and veins many straight, (semi-)appressed hairs,
distinctly two-coloured when dry. Petiole 2-7 cm
long. Stipules often persistent, orbicular in out-
line, digitately and deeply divided into 6-8 lobes,
the largest of those pinnatifid, 1-1.5 cm long,
hairy. Inflorescence panicle-like, a compound ra-
ceme with di- or monochasial last branches, up to
12 side-branches, the lower ones in the axils of
leaves, the entire thyrse up to 35 cm long, side-
branches up to 17 cm long. Bracts large, deeply
dentate. Pedicels l-2(-4) cm long, densely hairy
as all axes in the inflorescence. Flowers bisexual,
flower buds ovoid, pointed. Hypanthium cup-
shaped, (4-)5-7 mm across, densely hairy outside
with long, straight hairs hiding the smaller curly
ones. Sepals triangular, pointed, outer ones (4-)
6-8 by 3-6 mm, inner ones narrower, uncovered
margins with (2-)3-5 teeth, up to 1.5 mm long.
indumentum outside as hypanthium. Petals orbicu-
lar to elliptic, 3-7.5 by 2.5-5.5 mm, apex notch-
ed, white. Stamens 50-100, glabrous, filaments
up to 4 mm, anthers 0.5-0.8 mm long. Pistils
50-90, ovaries glabrous, on elevated, densely
hairy torus, style up to 5.5 mm long. Collective
fruits globular, up to 7 mm diam. when dry,
sepals closing after anthesis. Fruits curved, c. 2.5
by 1.5 mm when dry, glabrous, yellow to orange,
sometimes red, mesocarp juicy, a thin layer when
dry.
Distribution - Sumatra, Java, Lombok.
Habitat - Light forest and more open places like
thickets, forest edges, clearings, secondary bush,
and near craters. Montane, altitude (900-)1200-
2950 m.
Uses - The fruits seem to be delicious.
Notes - Some specimens from Java have only
female flowers.
The bullate leaves, the rather stout and coarse
habit, and the wide and lax inflorescences separate
the species rather clearly from the related R. moluc-
30. Rubus cumingii Kuntze, Meth. Species-
beschr. (1879) 72, 76; Merr., Enum. Philipp.
How. PI. 2 (1923) 227. — Type: Cuming s.n.,
Philippines.
Shrubs, dimensions unknown. Stems with
some vestiges of thin, curly hairs and also with
long and thicker hairs, prickles rather many, small.
Leaves ovate, not lobed, 7-8 by 5-6 cm, base
subtruncate, margin serrate, apex acute, herbaceous,
nervation pedate with 7 pairs of nerves, venation
transverse, both sides with long (semi-)appressed
hairs on the nerves and fewer on and between the
veins. Petiole 1.5 cm long. Stipules elliptic, en-
tire. Inflorescence panicle-shaped, c. 15 cm long
and wide, with 4 side branches, rich-flowered,
densely patently hairy. Bracts persistent. Pedicels
c. 3 mm. Flowers unisexual, flower buds ovoid.
Hypanthium cupular, c. 5.5 mm across, densely
hairy outside. Sepals broadly ovate, pointed, outer
ones 5 by 4 mm, inner ones narrower, uncovered
margins with one short tooth, indumentum outside
as hypanthium. Petals persistent, elliptic, 3.5 by 2
mm. Stamens not seen, staminodes in female
flowers c. 60, glabrous. Pistils 14, ovaries long-
hairy on the back near the apex, on little elevated,
hairy torus, style c. 2.5 mm long, hairy at base.
Collective fruits not known. Fruits c. 2.5 by 1.5
mm.
Distribution - Known from one duplicate of the
type, probably from Luzon, with female flowers.
272
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Note - Steiner 2027 (Luzon) and Be gain 1546
(Ternate) may be conspecific but differ in details of
the indumentum. Related to R. luzoniensis.
31. Rubus elongatus J.E.Smith, PL Icon.
Hact. Ined. 3 (1791) t. 62; Blume, Bijdr. (1826)
1112; Miq., Fl. Ind. Bat. I, 1 (1855) 380, incl.
varieties; Merr., Enum. Bom. Flow. PL (1921)
288; Ridley, Fl. Mai. Penins. 1 (1922) 679;
Backer & Bakh.f., Fl. Java 1 (1964) 516. —
Type: Commerson s.n., Java.
Rubus lobbianus Hook., Icon. PL 8 (1848) pi. 741/
742. — Rubus moluccanus L. var. lobbianus
(Hook.) Kuntze, Rev. Gen. PL 1 (1891) 222.
— Type: Lobb 62, Java.
Rubus blumei Focke, Bibl. Bot. 72 (1910) 60. —
Type: Korthals s.n., Java.
Rubus magnibracteatus Ridley, J. Fed. Malay St.
Mus. 8 (1917) 32. — Type: Robinson & Kloss
s.n., Sumatra.
Rubus elongatus J.E. Smith var. laevicalyx Rid-
ley, J. Fed. Malay St. Mus. 8 (1917) 31. —
Type: Robinson & Kloss 143, Sumatra.
Climbing, scrambling, or creeping shrubs.
Stems up to 25 m long, woody, thick, bark dark-
brown to black, densely hairy to almost glabrous,
prickles few to many, short, straight. Leaves
ovate, entire or shallowly lobed, 7.5-15 by 5-9.5
cm, base deeply cordate to truncate (in smaller
leaves), margins serrate to dentate, apex acute,
firmly herbaceous, nervation pedate with 8-11
pairs of nerves, venation transverse, densely hairy
on nerves and veins to almost glabrous above,
with a dense mat of short, curly hairs below, es-
pecially on nerves and veins covered by longer,
semi-appressed hairs, distinctly two-coloured in
living as well as in dried state. Petiole 2.5-5.5
cm long. Stipules early deciduous, suborbicular to
elliptic in outline, 6-9 by 6-10 mm, pinnatipar-
tite with 5-9 pairs of lobes, hairy outside. Inflo-
rescence a compound raceme, 12-35 cm long,
with 6-18 side branches, the lower ones in axils
of leaves, up to 10 cm long, laterals usually
branched again, with up to 25 flowers. Bracts
lobed. Pedicels up to 3 mm long, densely to slight-
ly hairy as are all axes, with 2 bracteoles halfway.
Flowers unisexual, plants dioecious, flower buds
globular. Hypanthiwn cup- to saucer-shaped, 3.5-5
mm across, densely tomentellous and with longer,
appressed hairs outside. Sepals triangular to (broad-
ly) ovate, 3-6 by 2.2-3.5 mm, apex acute or
rounded and shortly apiculate, margins entire, red
to purple as are hypanthium, pedicels, and bracts,
indumentum outside as hypanthium. Petals obcor
date to obovate, 5-11 by 4-6 mm, apex rounded
or emarginate, white to pink. Stamens 80-125,
glabrous, filaments 2-2.5 mm, anthers 0.8-1.2
mm long, staminodes 60-90, minute. Pistils
45-70, ovaries glabrous, on elevated, hairy torus,
style c. 2 mm long, pistillodes c. 1 mm. Collective
fruits globose, 6-10 mm diam when dry, sepals
closing after anthesis, spreading under ripe fruit.
Fruits sickle-shaped, 2-3.5 mm long, black when
ripe, mesocarp a rather thin layer when dry.
Distribution - Sumatra, Malaya, Borneo, W Java,
N Celebes, ? Moluccas (Tidore).
Habitat - Forests, thickets, near rivers and roads,
300-2300 m altitude.
Note - Field observations on the sex ratio are
wanting. Almost 80% of the existing herbarium
collections has female flowers. This may, how-
ever, not reflect the real situation but the inclina-
tion of collectors to pick fruiting specimens. See
also P. luzoniensis.
32. Rubus glomeratus Blume, Bijdr. (1826)
1111; Miq.,Fl. Ind. Bat. 1, 1 (1855) 381; Koord.,
Exk. Flora Java 2 (1912) 324. — Rubus mo-
luccanus L. var. glomeratus (Blume) Backer,
Schoolfl. Java (1911) 458; Backer & Bakh.f.,
Fl. Java 1 (1964) 517. — Non Hook, f., Fl.
Brit. India 2 (1878) 328 et auctt. al. — Type:
Blume s.n., Java (L, sheet nr 905.130-133/
134).
Rubus glabriusculus Hassk., Flora 27 (1844) 586;
Miq., Fl. Ind. Bat. I, 1 (1855) 383. — Rubus
moluccanus L. var. glabriusculus (Hassk.)
Kuntze, Rev. Gen. PL 1 (1891) 222. — Type:
not indicated.
Rubus sundaicus auct. non Blume: Kuntze, Meth.
Speciesbeschr. (1879) 60, 76.
Rubus ledermannii Focke, Bot. Jahrb. 56 (1916)
79; P. van Royen, Phan. Mon. 2 (1969) 91,
f. 23. — Type: Ledermann 11651 (lost), neo
van Royen NGF 30159, New Guinea.
Rubus robinsonii Ridley, J. Fed. Mai. St. Mus. 8
(1917) 31. — Type: Robinson & Kloss 132 (?),
Sumatra.
Rubus ledermannii Focke var. beleensis P. van
Royen, Phan. Mon. 2 (1969) 93, f. 24. — Type:
Brass 11035, New Guinea.
Climbing, trailing, or scrambling shrubs, up to
3 m high. Stems sparsely to densely hairy with
curly hairs never forming a closed felt, and with
longer, thicker, straight hairs, glabrate and then
dark brown to blackish, prickles usually few,
weak. Stipular cataphylls at the base of lateral
branches. Leaves broadly ovate, 4-15 by 3.5-13
Kalkman
Rosaceae
273
cm, shallowly 3(-5)-lobed or almost unlobed,
base cordate to (upper leaves) truncate, margins
serrate, apex acute, coriaceous, nervation pedate
with 5-7 pairs of nerves, venation reticulate, up-
per surface sometimes slightly bullate between
nerves and veins, hairy and glabrescent, lower sur-
face with on nerves and veins short, curly hairs
that never form a closed felt, and with longer and
thicker, straight hairs. Petiole 1— 5(— 7) cm long.
Stipules early deciduous, elliptic, 7-20 by 5-10
mm, dentate to pinnati partite with up to 6 pairs of
teeth or lobes, hairy outside. Inflorescence a termi-
nal thyrse with up to 6 lateral cymes of up to 3
flowers, up to 10 cm long, peduncle up to 5(-7)
cm. Bracts pinnatipartite. Pedicels up to 5(— 10)
mm long, hairy as are the other axes. Flowers bi-
sexual, flower buds ovoid, pointed. Hypanthium
cupular, 4-7 mm across, densely woolly outside
and with long, straight hairs, rarely with short,
stalked glands. Sepals triangular, outer ones 6-9
by 4-7 mm, inner ones narrower, apex sharply
pointed, not-covered margins with 2-6 teeth of
2-3 mm, covered margins entire, indumentum
outside as hypanthium. Petals early deciduous,
suborbicular to obovate or obcordate, 4.5-10 by
3-8 mm. apex rounded or emarginate, white,
sometimes hairy outside. Stamens 24-120, in
New Guinea up to 60, filaments up to 6 mm long,
glabrous, rarely hairy, anthers 0.5-1 mm long, gla-
brous or with 1-5 hairs on the top. Pistils 30-60,
ovaries glabrous, on elevated, hairy torus, style up
to 6 mm long. Collective fruits globular, up to 8
mm diam., sepals closing after anthesis and stay-
ing erect around ripe fruits. Fruits curved, 2-4 mm
long when dry, orange-red to red, mesocarp juicy, a
thin membranous layer when dry.
Distribution - Sumatra, Malaya, Java, S Cele-
bes, New Guinea, New Britain.
Habitat - Secondary and disturbed forests, and in
open places like clearings, riverbanks, landslides,
roadsides, forest edges, shrubland. Altitude (1200-)
1600-3000 m.
Notes - Specimens from Java and Sumatra dif-
fer slightly from the New Guinean specimens (/?.
ledermannii). Omitting overlapping measurements
of flower parts, the differences are as follows. Java
and Sumatra: stamens 75-120, anthers glabrous;
New Guinea: stamens 24-60, anthers usually
with 1— 2(— 5) hairs on top, rarely glabrous. The
material from Celebes resembles the West Male-
sian specimens. To recognize a separate species for
the New Guinean plants seems not to be justified.
The Malayan specimens are also slightly differ-
ent and were earlier recognized as var. gracilis King,
J. As. Soc. Beng. 66, ii (1897) 295.
Records from Moluccas and Philippines (/?. van-
overberghii Merr., var. pileanus Focke) could not
be substantiated but are not necessarily incorrect.
See Kalkman, Blumea 29 (1984) 380, note 5.
Rubus glomeratus has often been confused with
R. moluccanus and the two were united by some
authors. However, they can be separated without
difficulty by means of the leaf indumentum: the
felt of thin, short, curly hairs is in the former spe-
cies never entirely closed and the leaf surface remains
visible, which is not the case in R. moluccanus.
33. Rubus heterosepalus Merr., Philipp. J. Sc.
20 (1922) 387; Enum. Philipp. How. PI. 2
(1923) 228. — Type: Ramos & Edano BS
37609, Mt Polis.
Climbing or straggling shrubs. Stems densely
hairy with many long, straight hairs and few short,
thin, curly ones, prickles up to 2 mm. Leaves
broadly ovate, 8.5-14 by 6.5-10 cm, shallowly
3-5-lobed, base cordate, margins rather grossly
serrate, apex gradually acuminate, coriaceous, ner-
vation pedate with 5-6 pairs of nerves, venation
reticulate, upper surface bullate between the veins,
both surfaces with long hairs on nerves and veins
and lower surface also with a dense felt of thin
curly hairs all over, two-coloured when dry. Petiole
3-5 cm long. Stipules pinnatipartite with 6-8
pairs of lobes, c. 15 by 11 mm, hairy outside. In-
florescence a compound raceme, large and rich,
10-25 cm long, only the lowermost of the 4-6
side branches in the axil of a leaf or all laterals in
the axils of bracts. Bracts persistent, pinnatisect,
large. Pedicels up to c. 8 mm long, densely hairy
as are the other axes. Flowers bisexual, flower
buds ovoid. Hypanthium cup-shaped, c. 8 mm
across, densely hairy outside. Sepals distincdy di-
morphous, outer ones 10— 14(— 18) by 10-13 mm,
with 5 or more pairs of lobes, those up to 7 by
1.5 mm, inner sepals triangular and not lobed,
10-11 by c. 4 mm, indumentum outside as hy-
panthium. Petals falling early, c. 6 by 3-3.5 mm,
apex deeply emarginate. Stamens 70-90, fila-
ments up to 4 mm, glabrous, anthers c. 0.8 mm
long. Pistils many, ovaries glabrous, on hairy
torus, style up to 4.5 mm long. Collective fruits
probably globular, c. 1 cm diam. (dry), sepals clos-
ing after anthesis, widely spreading under ripe
fruits. Fruits 3 mm long, mesocarp rather thick.
Distribution - Luzon.
Habitat - Mossy forest at 1800 m, according to
Merrill (I.e., 1922).
Note - Only few specimens known. Related to R.
chrysophyllus, but sepals larger, with longer lobes.
274
Flora Malesiana ser. I, Vol. 11 (2) (1993)
34. Rubus keleterios P. van Royen, Phan.
Monogr. 2 (1969) 87, f. 22. — Type: Hoogland
& Pullen 6226, Mt Hagen.
Shrubs. Stems densely hairy, prickles few,
short. Leaves broadly ovate in outline, 5-7.5 by
5-6.5 cm, distincUy 3-lobed, base cordate, margins
unevenly serrate, apex acute, firmly herbaceous,
nervation palmate with 5 main nerves, venation
reticulate, rather densely hairy above, lower surface
with a dense woolly felt of short, thin, curly hairs,
and with many thicker, straight, long hairs, dis-
tinctly two-coloured. Petiole 1.5-2.5 cm long.
Stipules rather persistent, linear, 14-17 by 2 mm,
with 2-3 narrow lobes, hairy outside. Inflores-
cence a leafy thyrse, c. 15 cm long, with c. 8 di-
chasial to monochasial laterals of c. 4 cm long,
with up to 5 flowers. Bracts c. 10 mm long, deep-
ly incised, bracteoles 2, on the pedicel, persistent
as are the bracts. Pedicels up to 1.5 cm long, den-
sely hairy as are the other axes. Flowers bisexual,
flower buds ovoid. Hypanthium cup-shaped, 7 mm
across, short-woolly and with longer hairs outside.
Sepals triangular, 6.5-8 by 2.5-4 mm, outer
ones usually with up to 3 pairs of teeth (up to 1
mm long), inner ones entire, indumentum outside
as hypanthium. Petals persistent, ± orbicular, dis-
tinctly clawed, 5.5-7 by 3-5 mm, white. Sta-
mens 85-125, filaments up to 5 mm, glabrous,
anthers c. 0.5 mm long. Pistils c. 60, ovaries gla-
brous or with few hairs on top, on elevated, dense-
ly hairy torus, style up to 6 mm long. Collective
fruits globular, c. 1 cm diam., sepals closing after
anthesis. Fruits c. 4 mm long (dry), exocarp gla-
brous or with long hairs on backside in the upper
half, mesocarp thick and leathery when dry, prob-
ably thick and juicy when living.
Distribution - New Guinea (only known from
two collections from Papua New Guinea); a col-
lection from Cook Distr., Queensland, Australia
(Clarkson 2742) seems to be conspecific.
Habitat - Once in shrubland, once in grass
swamp, 1700-1950 m alt The Queensland collec-
tion is from 780 m altitude.
Note - Closely related to R. moluccanus but
distinct by its leafshape (very broad and short, end
lobe about half of total leaf length) and its long
pedicels with persistent bracteoles.
35. Rubus luzoniensis Merr., Philipp. J. Sc,
Suppl. 1 (1906) 195; Elmer, Leafl. Philipp. BoL
2 (1908) 452; Merr., Enum. Philipp. Row.
PI. 2 (1923) 228. — Type: Merrill 4596, Mt
Data.
Climbing or sprawling shrubs, gynodioecious?
Stems up to 10 m long, densely hairy when young,
prickles rather few to many, short but strong.
Leaves (broadly) ovate to suborbicular, 5-8.5 by
4-6.5 cm, not or very shallowly lobed, base shal-
lowly cordate or truncate, margins serrate, apex ob-
tuse to acute, stiff -coriaceous, nervation pedate with
5-7 pairs of nerves, venation transverse, (rather)
densely hairy above, lower surface densely short-
woolly all over and with long, straight hairs on
nerves and veins, distinctly two-coloured. Petiole
1-3 cm long. Stipules often rather persistent, lin-
ear, 12-19 by 3-4 mm, entire or minutely teeth-
ed, hairy outside. Inflorescence a panicle-like com-
pound raceme, richly and widely branched and
many-flowered, up to 40 cm long, up to 15 side
branches, up to 18 cm long. Bracts rather large,
persistent. Pedicels up to 5 mm long, densely
hairy as all axes in the inflorescence. Flowers bi-
sexual or female. Hypanthium saucer-shaped, 3-
4.5 mm across, densely hairy outside. Sepals ovate,
5-6 by 2.5-4 mm, after anthesis growing to 8
by 6 mm, apex apiculate, margins entire or with
minute teeth, on the outside with a woolly felt and
the not-covered parts also with long appressed hairs.
Petals obovate to elliptic, 5-6.5 by 2.5-4 mm,
apex obtuse to emarginate, white. Stamens 50-
100, filaments up to 4.5 mm, glabrous, anthers
0.5 mm long, staminodes with shorter filament
and minute anther. Pistils 15-25, ovaries long
hairy at the top on the backside, on a slightly ele-
vated, long-hairy torus, style 2-3 mm long, some
hairs at the base. Collective fruits consisting of
few fruits only, sepals closing after anthesis. Fruits
c. 3 mm long when dry, still hairy at apex, bright
red, mesocarp a rather thick, tough layer when dry.
Distribution - Luzon.
Habitat - Forest and clearings, altitude 2000-
2500 m.
Notes - The sex distribution is incompletely
known. The great majority of the rather few herba-
rium collections have female flowers. The species
is most closely related to R. elongatus and replaces
that species in the Philippines.
36. Rubus malvaceus Focke, Bibl. Bot. 72
(1910) 81, f. 30; Thuan, Fl. Camb., Laos &
Vietnam 7 (1968) 51, excl. specimens from
Asian continent; Fl. Thailand 2 (1970) 56, idem.
— Rubus moluccanus L. var. malvaceus (Focke)
Backer, Schoolfl. Java (1911) 458; Backer &
Bakh. f., Fl. Java 1 (1964) 517. — Type: Ploem
19980, Sindanglaya, W Java.
Kalkman — Rosaceae
275
Rubus wichurae Focke, BIbl. Bot. 72 (1910) 79.
— Type Wichura 2092, Mt Tangkubanperahu,
W Java.
Shrubs with overhanging branches. Stems with
a dense, long persistent indumentum, prickles not
many, curved, up to 3 mm. Leaves broadly ovate
to suborbicular in outline, 7— 15(— 19) by 7— 15(— 17)
cm, distincUy 5- to 7-lobed, base deeply and sharp-
ly cordate, margin rather evenly serrate, apex usu-
ally acute, coriaceous, nervation 5-palmate, the
lowermost main nerves pedately nerved, venation
reticulate, nerves and veins impressed above and
surface distinctly bullate between them, densely
hairy above, lower surface with a dense felt of long,
curly hairs all over and with many long, patent
hairs on nerves and veins, distinctly two-coloured.
Petiole 1.5-6(-8) cm long. Stipules rather per-
sistent, pinnatipartite to digitate-pinnate, 15-18
by 12-18 mm, the lobes 0.5-1.5 mm wide, hairy
outside and on margins. Inflorescence a large thyrse,
20-25 cm long, with up to 12 cymose laterals in
the axils of normal or smaller leaves or bracts, the
laterals up to 3 cm long, with 2-4(-10) flowers.
Bracts pinnatifid, large. Pedicels up to 1 cm long,
hairy. Flowers bisexual, the flower buds ovoid.
Hypanthium cupular, 6-10 mm across, densely
hairy outside. Sepals recurved at the top during
anthesis, triangular, 7-10 mm long, outer ones
4-8 mm wide, the not-covered margins long-
dentate, inner ones 3.5-6 mm wide, entire, in-
dumentum outside as hypanthium. Petals sub-
orbicular, 5-6 by 5-5.5 mm, clawed, the mar-
gin undulating, white. Stamens 100-200, fila-
ments up to 5 mm, glabrous, anthers 0.5-0.8
mm long, with long hairs on top. Pistils 60-80,
ovary glabrous, the style up to 7.5 mm long. Col-
lective fruits not seen. Fruits 2-3 mm long, the
colour unknown.
Distribution - W Java, Sumba.
Habitat - Hardly any data, one collection from
1300 m altitude.
Notes - Quite closely related to R. rugosus J.E.
Smith, which is known from the Asian continent
and Sri Lanka. It seems, however, premature, to
combine the two.
The patchy distribution cannot be explained by
supposing an escape from the botanical garden at
Cibodas, Java.
37. Rubus mearnsii Elmer, Leafl. Philipp. Bot.
2 (1908) 448; Merr., Philipp, J. Sc, Bot. 5
(1910) 353; Enum. Philipp. How. PI. 2 (1923)
228. — Type: Mearns BS 4304, Luzon.
Shrubs. Stems rather densely hairy when young
and with many c. 1 mm long gland-bearing hairs,
prickles (rather) many, curved, up to 1.5 mm.
Leaves broadly ovate to suborbicular, 5-9 by 5-7
cm, not or faindy lobed, base cordate, margin rather
coarsely serrate, apex obtuse to acute, coriaceous
to herbaceous, nervation pedate with 6-7 pairs of
nerves, venation transverse, both surfaces rather
sparsely hairy, more densely on midrib and nerves
below. Petiole 1-2 cm long. Stipules persistent,
tongue-shaped, 7-9 by 3-4 mm, with 5-6 pairs
of small teeth, hairy and glandular outside. Inflo-
rescence a thyrse, laxly paniculate in appearance,
up to 52 cm long, with up to 18 lateral branches
of up to 24 cm long and with up to 30 flowers.
Bracts up to 9 mm long, hairy and glandular out-
side. Pedicels 0.5-1.5 cm long, long-hairy and
with glandular hairs, as are the other axes. Flowers
bisexual, flower buds ovoid, pointed. Hypanthium
saucer-shaped, 6 mm across, hairy and glandular-
hairy outside. Sepals ovate, 7-8 mm long, outer
ones 5-6 mm wide, with 3-6 teeth on the un-
covered margins, inner ones c. 4 mm wide, entire,
indumentum outside as hypanthium. Petals none.
Stamens 60-90, filaments up to 5 mm, glabrous,
anthers 0.5-0.8 mm long. Pistils 12-18, ovaries
glabrous, on flat, hairy torus, style up to 4 mm
long. Collective fruits c. 6 mm diam., loose, often
only some of the fruits developing, sepals closing
and remaining closed after anthesis. Fruits 4 mm
long when dry, mesocarp fleshy, a rather tough
layer when dry.
Distribution - Luzon.
Habitat - Only six collections investigated, the
majority without field data. According to Merrill
(1923) in mossy forest, c. 2400 m altitude, one
specimen collected along the road at c. 2260 m.
Note - Closely related to R. benguetensis.
38. Rubus moluccanus L., Spec. PI. (1753)
1197; Blume, Bijdr. (1826) 1109; Miq., Fl.
Ind. Bat. I, 1 (1855) 382; Backer & Bakh.f.,
Fl. Java 1 (1964) 516, excl. var. glomeratus
(Blume) Backer and var. malvaceus (Focke)
Backer; P. van Royen, Phan. Monogr. 2 (1969)
98, incl. var. austropacificus P. van Royen and
var. thespesiaephyllos P. van Royen. — Type:
Rumphius, Herb. Amboin. 5 (1747) 88, pi. 47,
f. 2 [Rubus moluccus latifolius].
Rubus sundaicus Blume var. discolor Blume, Bijdr.
(1826) 1111. — Rubus moluccanus L. var. dis-
color (Blume) Kalkman, Blumea 29 (1984) 359.
— Type: Kuhl & van Hasselt s.n., Java.
276
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Rubus hasskarlii Miq., Fl. Ind. Bat. I, 1 (1855)
381. — Rubus moluccanus L. var. hasskarlii
(Miq.) Kuntze, Rev. Gen. PI. 1 (1891) 222. —
Type: not identified, Java.
Rubus moluccanus L. var. obtusangulus Miq., Fl.
Ind. Bat. I, 1 (1855) 383, 'obtusangula'; Kalk-
man, Blumea 29 (1984) 362. — Type: Junghuhn
s.n., Java.
Rubus angulosus Focke, Bibl. BoL 72 (1910) 90,
f. 35, nom. illeg., non Gremli (1871); Koord.,
Exk. Fl. Java 2 (1912) 324; Ridley, Fl. Mai.
Penins. 1 (1922) 678, f. 59. — Rubus moluc-
canus L. var. angulosus Kalkman, Blumea 29
(1984) 364 — Kurz (Amann) s.n., Bangka,
lecto.
Rubus hasskarlii Miq. subsp. dendrocharis Focke,
Bibl. Bot. 72 (1910) 99, f. 42. — Rubus den-
drocharis (Focke) Focke, Bot. Jahrb. 54 (1916)
70. — Rubus moluccanus L. var. dendrocharis
(Focke) P. van Royen, Phan. Monogr. 2 (1969)
106, f. 28. — Type: Rodatz & Klink 182, New
Guinea, in SING.
Rubus glomeratus auct. non Blume: Ridley, Fl.
Mai. Penins. 1 (1922) 679.
For a more complete synonymy, see Kalkman,
Blumea 29 (1984) 346.
Climbing or scrambling, rarely creeping shrubs.
Stems up to 6(-10) m long, densely hairy when
young, (tardily) glabrescent, prickles usually not
many, small. Leaves ovate to broadly ovate in out-
line, 6-20 by 4-15 cm, variously lobed, base
cordate to subtruncate, margins serrate, apex acute
to acuminate, (firm)-herbaceous, nervation pedate
with 5-9 pairs of nerves or palmate with 7 main
nerves (var. angulosus), venation (widely) reticulate,
the surface between the veins not or indistinctly
raised above, upper surface hairy, especially on the
nerves, lower surface with a densely woven felt of
long, thin, curly hairs all over, and with on nerves
and veins usually many long, thicker, straight, ap-
pressed to patent hairs, distincdy two-coloured.
Petiole 2-6 cm long. Stipules early falling, 7-17
by 4-12 mm, pinnatilobed to pinnatipartite with
4-10 pairs of lobes, sometimes digitate, lobes up
to 8 by 1 mm, hairy outside. Inflorescence a ter-
minal, leafy, compound raceme, up to 20(-50) cm
long, with up to 12 laterals, those up to 5(-9) cm
long and with up to 10(-30) flowers. Bracts up to
17 by 9 mm. Pedicels up to 1 cm long, hairy as
are all axes. Flowers bisexual, flower buds ovoid,
pointed. Hypanthium cupular, 4-7 mm across,
densely woolly and with patent to appressed straight
hairs outside. Sepals erect or apically recurved in
anthesis, triangular to ovate, 4-9 by 2-6 mm,
apex acute to pointed, not-covered margins of the
outer sepals with one or few, up to 3 mm long
teeth, covered margins entire, indumentum outside
as hypanthium. Petals long remaining, suborbicu-
lar to elliptic, 3-7 by 3-6 mm, apex rounded to
emarginate, white, rarely reported to be pink, red,
or yellow. Stamens 30-185, filaments up to c. 4
mm, anthers 0.2-0.7 mm long, mostly with few
to several long hairs on top or on connective. Pis-
tils 30-135, ovaries glabrous, on elevated, hairy
or glabrous torus, styles up to 9 mm long. Collec-
tive fruits globular, 0.8-1 cm diam. when dry,
sepals closing or apically slightly recurved after
anthesis, spreading at ripeness. Fruits 2-3 by 1-2
mm when dry, red, mesocarp thick and fleshy, only
a thin layer when dry. - Figs. 5, 6.
Distribution - All over Malesia, extending to
the North to Sri Lanka (introduced?), Thailand, and
Vietnam, to the South and East to Queensland,
Carolines, New Hebrides, Fiji, and New Caledonia.
Habitat - Essentially at low and medium alti-
tudes, up to 2000 m, occasionally higher.
Uses - Information from herbarium labels is re-
ported under the varieties. According to literature
[Burkill, Diet. Econ. Prod. Mai. Penins. (1966)
1952; Heyne, Nutt. PI. Indon. (1950) 693; Quis-
umbing, Medic. PI. Philipp. (1951) 354; Woodley
(ed.), Medic. PI. Papua New Guinea I, Morobe
Prov. (1991) 120] the species has a number of wide-
spread medicinal uses. The sap of shoots, the
chewed leaves or decoctions of roots are obviously
effective in relieving internal pains, in the treat-
ment of dysentery or diarrhoea, sprue and angina,
and for external afflictions like sores and boils.
Several times the use as an emmenagogue (stimu-
lating menstruation) or abortifacient is mentioned,
but also applications to prevent miscarriage, which
seems a strange combination. Use of the fruits as a
remedy for children's bed-wetting was mentioned
by Rumphius and later often repeated. It is possible
that the medicinal uses recorded for this species
also are valid for other species of the genus.
Notes - Rubus moluccanus L. is taken here in
the narrowest possible sense. See Kalkman, Blumea
29 (1984) 349, for a discussion on the species
limits as observed by Kuntze [Mcth. Speciesbeschr.
(1879) 33, and Rev. Gen. PI. 1 (1891) 222] and
Focke [Bibl. Bot. 72 (1910) 88]. Rubus molucca-
nus has been considered by Backer [Schoolfl. Java
(191 1) 457, and later] and by others as a taxonom-
ical problem of the same order as presented by R.
fruticosus, but incorrectly so. Rubus moluccanus
is, if properly delimited from neighbouring but
recognizable species, not much more variable than
any other widely distributed species. There is no
Kalkman — Rosaceae
277
Fig. 5. Rubus moluccanus L. Leaf shapes of the varieties: a, b. var. moluccanus; c. var. obtusangulus Miq.
d, e. var. discolor (Blume) Kalkman; f. var. angulosus Kalkman.
evidence at present for supposing that the species
is apogamous.
The species was divided by Kalkman into four
varieties, based on leaf characters. Intermediates do
occur and inadequately collected material showing
- as too often is the case - only the upper leaves
of lateral branches, cannot always be identified
down to the variety.
KEY TO THE VARIETIES
la Leaf-base subtruncate to shallowly cordate, the
basal lobes making an obtuse angle. The
straight hairs on nerves and veins below ap-
pressed d. var. obtusangulus
b. Leaf-base cordate, the angle between the mar-
gins of the basal lobes sharp 2
278
Flora Malesiana ser. I, Vol. 11 (2) (1993)
2a. The straight hairs on nerves and veins below
appressed to semi-appressed
b. var. discolor
b. The straight hairs on nerves and veins below
patent 3
3a. The basal lobes of the leaves distincdy over-
lapping or at least touching each other. Leaves
3_5(_7)-lobed, the lateral incisions up to 2 cm
deep, the nervation 7-palmate to -pedate
a. var. angulosus
b. The basal lobes of the leaves not overlapping,
their margins parallel or making a sharp angle.
Leaves not lobed or shallowly 3-5-lobed, the
lateral incisions 0.5-1 cm deep, the nervation
5- or 7-pedate c. var. moluccanus
a. var. angulosus Kalkman, Blumea 29 (1984)
364. See there for synonymy.
Young twigs and nerves and veins on the lower
leaf surface with semi-appressed to patent, long,
straight hairs over the woolly felt of thin, curly
hairs. Leaves distinctly 3-5(-7)-lobed with the
main lobes usually shallowly lobed again, apical
lobe large (half of the total leaf length or slightly
smaller), length/width index (1.1-)1.2-1.3(-1.4),
base cordate, the margins of the basal lobes dis-
tinctly overlapping or at least touching each other.
Stamens (85-)100-185, anthers hairy on apex. -
Fig. 5f.
Distribution - Thailand, Vietnam, Andaman Is.,
Nicobar Is.; Malesia: Sumatra and islands near it,
Malaya, Singapore, Java, Borneo, Palawan, Cele-
bes, Lesser Sunda Islands.
Habitat - Forest edges, secondary forest, thickets,
also near the beach and on riverbanks, altitude 0-
500(-1000) m, the few specimens seen from Java
from higher altitude (up to 1450 m).
Uses - Fruits edible. Boiled roots are a medicine
against dysentery (Malaya, Alvins 375).
b. var. discolor (Blume) Kalkman, Blumea 29
(1984) 359. See there for synonymy.
Young twigs and nerves and veins on lower
leaf-surface with appressed (rarely semi-appressed)
straight hairs over the woolly cover of curly hairs.
Leaves shallowly 3-5-lobed, sometimes hardly
lobed, rarely more distinctly lobed, length/width
index (1.0-)1.2-1.5(-2.5), basal incision narrow
with subparallel margins or a sharp angle between
them, the leaves in and close under the inflorescence
often with a wider angle or base even subtruncate.
Stamens 70-180, anthers hairy at apex or glabrous.
-Fig. 5d, e.
Distribution - Malesia: Sumatra, Malaya (Pa-
hang, Penang I., and Tioman I.), Borneo, Java,
Philippines, Celebes incl. Buton I., Lesser Sunda
Islands, Moluccas, New Guinea, Bismarck Archi-
pelago. Also in Solomon Islands, New Hebrides
(Erromanga), and New Caledonia.
Habitat - Secondary and shrubby vegetation, in
primary forest especially in clearings, along paths
and in the forest margin, but possibly also in closed
forest, further on hillsides, roadsides, riversides,
from sea-level up to 2000 m altitude, highest re-
cords c. 2500 m and (once) 2900 m.
Uses - The fruits are edible but tasteless. The
fluid from the stems is applied for eye diseases (Java,
KM & van Hasselt), fluid from the leaves is ap-
plied to the eyes of young mothers (Borneo, Winckel
1013). Young twig ends eaten raw as vegetable
(Java, Bakhuizen van den Brink 8168).
Notes - The longer leaves with leaf index > 1.5
are found especially in Sumatra and Malaya (up to
2.0) and in New Guinea (up to 2.5).
Two specimens from Mt Panggrango on Java
(Schiffner 2019, van Steenis 17620) have peculiar
gland-bearing bristles on vegetative parts and on
the hypanthium and sepals.
For distinction with var. obtusangulus, see there.
c. var. moluccanus — For synonymy, see Kalk-
man, Blumea 29 (1984) 346.
Young twigs and nerves and veins on the lower
leaf-surface with (apart from the underlying woolly
felt) patent to semi-patent, long, straight hairs.
Leaves shallowly 3-5-lobed or not lobed at all,
length/width index (1.0-)1.1-1.4(-1.8), basal in-
cision narrow with parallel margins or a sharp
angle between them. Inflorescence usually leafy
but sometimes all lateral racemes in axils of bracts.
Stamens 30-1 50(- 200), anthers hairy at apex or
glabrous. - Fig. 5 a, b.
Distribution - Sri Lanka [introduced according
to Tirvengadum, Fl. Ceyl. 3 (1981) 353], Thailand,
Vietnam; Malesia: Sumatra, Malaya incl. Penang
I. and Langkawi I., Singapore, Borneo, Java, Phi-
lippines, Celebes, Lesser Sunda Islands, Moluccas,
New Guinea, New Britain. Also in the Solomon
Islands, Carolines, Fiji Islands, and Australia
(Queensland).
Habitat - Forest edges, secondary forest, lighter
places in primary forest (e.g. riverbanks), open
places like thickets, roadsides, lavastreams, heath
vegetation, from sea-level up to c. 2000 m, very
rarely higher.
Uses - The fruits are edible but have a poor fla-
vour according to several collectors. Young, fresh
Kalkman — Rosaceae
279
■fyei sen doi-p 03
1 cm
Fig. 6. Rubus moluccanus L. var. obtusangulus Miq. a. Flowering branch; b. old flower (a: Schodde 1493;
b: Streimann NGF 34081).
280
Flora Malesiana ser. I, Vol. 11 (2) (1993)
leaves are a cure for diarrhoea (Borneo, Mamit S
33363). Twigs may be used for bundling firewood
(Philippines, Conklin & Buwaya PNH 78635) and
are used for catching bats (Philippines, Britton 434,
how is not described).
d. var. obtusangulus Miq., Fl. Ind. Bat. 1, 1
(1855) 383; Kalkman, Blumea 29 (1984) 362.
See there for synonymy.
Young twigs and nerves and veins on the lower
leaf surface with appressed (sometimes semi-ap-
pressed), long, straight hairs over the woolly layer
of curly hairs, the straight hairs sometimes only
sparsely present. Leaves shallowly 3-5-lobed to
hardly lobed at all, length/width index (1.1— )1.3—
1.7(— 1.9), base subtruncate to shallowly cordate
with an obtuse angle between the margins of the
basal lobes. Stamens 30-140, anthers nearly al-
ways hairy on top, but in Sumatra always glabrous.
-Figs. 5c, 6.
Distribution - Burma, Thailand, Vietnam; Ma-
lesia: Sumatra, Malaya, Borneo, Java, Philippines,
New Guinea, New Britain.
Habitat - Forest edges, secondary forest, thickets,
along roads, trails, and streams, altitude (150-)9OO-
2000(-3000) m.
Uses - The fruits are edible. Stems are used as
binding material (Philippines, Conklin & Buwaya
PNH 80574). Men wash their bodies with the leaves
before going to fight (Papua New Guinea, Maprik
Subprov., Wiakabu LAE 73551).
Note - The recognition in the herbarium of the
varieties discolor and obtusangulus does usually
not present a problem, if the specimen possesses a
number of leaves below the inflorescence. If only
the flowering part of a twig is present, identifica-
tion is not reliably possible since the raceme-bear-
ing leaves of var. discolor are often subtruncate at
the base.
39. Rubus perfulvus Merr., Philipp. J. Sc. 20
(1922) 386; Enum. Philipp. Flow. PI. 2 (1923)
229. — Type: Ramos & Edaho BS 38566, Min-
danao.
Climbing shrubs. Stems densely hairy when
young, prickles few, short. Leaves ovate, 5-9 by
4-7 cm, not or hardly lobed, base truncate to
rounded, margin grossly dentate, apex acuminate,
stiff coriaceous, 3-nerved, the two lateral main
nerves with c. 4 basiscopic side-nerves, the middle
nerve with c. 4 pairs of lateral nerves, nerves and
veins impressed above and leaf surface bullate, up-
per surface scattered long-hairy, glabrescent, lower
surface with a dense and thick woolly felt of curly
hairs and with longer, straight, appressed hairs on
nerves and veins, distinctly two-coloured. Petiole
1-1.5 cm long. Stipules persistent, up to 15 by 3
mm, entire. Inflorescence according to Merrill a
terminal, 5-flowered, compact raceme and also
some solitary flowers in the upper leaf axils.
Flowers large. Sepals triangular, c. 12 by 3-4 mm,
outer ones with few marginal teeth. Petals not
seen. Stamens probably with glabrous anthers.
Collective fruits c. 1.2 cm diameter. Fruits gla-
brous (Merrill), red, the mesocarp a thin mem-
branous layer when dry, endocarp rugose, stone
c. 3 mm long.
Distribution - Only known from the type speci-
men from Mt Lipa.
Habitat - Mossy forest, c. 2000 m altitude.
Note - Insufficiently known, complete inflores-
cences and flowers in anthesis not seen. Relation-
ship is probably closest to R. rolfei Vidal.
40. Rubus pyrifolius J.E.Smith, PI. Icon.
Hact. Ined. 3 (1791) t. 61; Miq., Fl. Ind. Bat. I,
1 (1855) 384; Merr., Enum. Philipp. How. PI.
2 (1923) 229; Backer & Bakh. f., Fl. Java 1
(1964) 516; Steenis, Mount. Fl. Java (1972)
pi. 45-6. — Dalibarda pyrifolia (J.E. Smith)
Blume, Bijdr. (1826) 1112. —Rubus moluc-
canus L. var. pyrifolius (J.E. Smith) Kuntze,
Rev. Gen. PI. 1 (1891) 222. — Type: Commer-
son s.n., Java.
Dalibarda latifolia Blume, Bijdr. (1826) 1112. —
Type: not found, from Cianjur, W Java.
Rubus philippinensis Focke in Elmer & Focke,
Leafl. Philipp. BoL 5 (1913) 1617; Merr., Enum.
Philipp. Row. PI. 2 (1923) 229. — Type: Elmer
13606, Mindanao.
For a more complete synonymy, see Kalkman,
Blumea 29 (1984) 333.
Climbing shrubs. Stems up to 8(-30) m long,
usually with few hairs, prickles usually few,
small, minute capitate brown hairs often present
on vegetative parts and inflorescences. Leaves el-
liptic to ovate, 6.5-16 by (2.5-)3.5-8.5(-9.5)
cm, not lobed, base rounded, margin serrate-cre-
nate, apex acute or obtuse and then shortly or long
acuminate, herbaceous, nervation pinnate with 5-8
pairs of nerves, the lowermost of these with up
to 6 strong, basiscopic side nerves, rarely truly
pedate, secondary nerves not reaching the margin,
venation widely transverse, usually rather densely
hairy on midrib and larger nerves above, lower
surface with (semi-)appressed hairs on nerves and
veins. Petiole 0.5-1 (-3) cm long. Stipules early
Kalkman
Rosaceae
281
falling, 6-9 by 0.5-1.2 mm, apically 3-6-lobed,
lobes up to 6 mm, hairy. Inflorescence broadly
paniculate, a leafy compound raceme, up to 30
(-40) cm, with 9-16 primary laterals, the latter
up to 12(-15) cm, with up to 10 side branches,
usually being dichasia with up to 7 flowers. Bracts
up to 9 by 1.5-2 mm, lobed. Pedicels 6-8(-10)
mm long, densely hairy as are the other axes in the
inflorescence. Flowers bisexual, flower buds ovoid,
pointed. Hypanthium saucer- to cup-shaped, 3-5
mm across, shortly woolly outside, sometimes
also with appressed, straight hairs. Sepals triangu-
lar to ovate, reflexed to horizontal during anthesis,
4— 7(— 1 1) by 2-4 mm, outer margins with 1 or 2
teeth or lobes of up to 2 mm, inner margins entire,
indumentum outside as hypanthium. Petals elliptic
to obovate, 2-5(-7) by 0.5-2(-3.5) mm, apex
(sub)truncate, usually sinuate or notched, white.
Stamens (40-)50-80(-100), filaments up to 6.5
mm, anthers 0.5-0.8 mm long, usually wider
than long, violet. Pistils (3— )5 — 10(— 17), ovaries
usually with many long hairs on the dorsal side,
rarely glabrous, on a low, long-hairy torus, styles
up to 5.5(-7) mm long, rarely with some hairs in
basal part. Collective fruits loose, usually less than
half of the ovaries developing, c. 0.5 cm diam.,
sepals closing after anthesis. Fruits 3.5-4.5 mm
long when dry, with some hairs, red, mesocarp
rather thick and fleshy.
Distribution - Laos, S Vietnam, Thailand, Chi-
na; Malesia: Sumatra, Borneo, Java, Philippines
(Negros, Leyte, Mindanao), Celebes, Lesser Sunda
Islands (Bali, Lombok, Flores).
Habitat - Forest and forest edges, altitude 500-
1700(-2200) m.
Ecology - Characteristic leaf galls, caused by a
gall-mite, are found in specimens from Java and
Sumatra. The galls are hollow bladders on the upper
leaf surface, purple coloured according to Docters
van Leeuwen & Docters Van Leeuwen-Reynvaan,
Zoocecidia of Neth. Indies (1926) 219. On the
underside of the leaf the bladders are densely hairy.
41. Rubus rolfei Vidal, Phan. Cuming. (1885)
171; Elmer, Leafl. Philipp. Bot. 2 (1908) 454;
Merr., Enum. Philipp. Flow. PI. 2 (1923) 230.
— Types: Cuming 808, lecto; Vidal 294; both
Luzon.
Rubus elmeri Focke, Bibl. Bot. 72 (1910) 112;
Merr., Enum. Philipp. Flow. PI. 2 (1923) 227.
— Types: Merrill BS 4651, Mearns BS 4305,
Luzon.
Rubus calycinoides Hayata, Icon. PI. Form. 3
(1913) 88, non Kuntze (1879). — Type: Mori
& Kato s.n., Taiwan.
Sprawling shrubs with prostrate long shoots up
to 1 or more m, and with axillary, ascending, leafy
shoots up to 25 cm long, terminating in an inflo-
rescence, more rarely climbing shrubs. Stems den-
sely hairy when young, prickles (very) few, small.
Leaves orbicular to triangular in oudine, 2-5.5
(-9) by 2-5(-9) cm, shallowly 3- to 5-lobed,
base cordate, basal lobes rounded, margin unevenly
crenate, apex rounded to acute, stiff -coriaceous, ner-
vation pedate, with 4-5 pairs of nerves, venation
widely reticulate, upper surface bullate between
nerves and veins, hairy but soon glabrous above,
lower surface with a dense woolly felt of thin,
curly hairs, and with on nerves and veins longer,
patent, straight hairs, distinctly two-coloured.
Petiole 0.5-3(-4.5) cm long. Stipules rather
long persistent, 6-10(-13) by 2.5-4(-7) mm,
incised. Inflorescence a terminal, simple raceme,
usually very dense, up to 3 cm long, with up to 8
flowers. Bracts up to 13 by 10 mm, incised. Pedi-
cels 0-5 mm long, densely hairy as is the rachis
of the raceme. Flowers bisexual, flower buds ovoid.
Hypanthium saucer-shaped, 4-6 mm across, den-
sely woolly and with straight, longer hairs outside.
Sepals triangular, outer ones 6-10 by 3-8 mm,
inner ones narrower, growing after anthesis, apex
acute to acuminate, uncovered margins with 1-3
small teeth, up to 3 mm, indumentum outside as
hypanthium. Petals orbicular with rounded apex,
up to 14 by 12 mm, white. Stamens 120-140,
filaments up to 5 mm, anthers 0.5-1 mm long,
glabrous or with few long hairs. Pistils 40-80,
ovaries glabrous, on elevated, hairy torus, styles
up to 5(-8) mm long. Collective fruits globular,
c. 1 cm, sepals closing after anthesis, spreading at
ripeness. Fruits 2-2.5 mm long when dry, yellow
to orange (also reported as pink or red), mesocarp
soft and thick, only a thin layer when dry.
Distribution - Taiwan; Malesia: Philippines
(Luzon, Negros, Mindoro).
Habitat - Forest and more open places like clear-
ings at higher altitudes, alt (900 -) 1500 -2700 m.
Uses - The fruits are edible and sweet.
Note - Most of the plants collected had prostrate,
woody twigs with erect, determinate, short laterals
terminating in an inflorescence [see Kalkman, Blu-
mea 29 (1984) 371, f. 5]. The last-but-one order
branches, however, are not always creeping but
sometimes climbing and maybe there are even
bushy plants with more or less erect branches.
Possibly this will be largely determined by local
conditions. Herbarium specimens of creeping
plants may look rather different from collections
from climbing shrubs, but in essential characters
except habit they are alike.
282
Flora Malesiana ser. I, Vol. 11 (2) (1993)
42. Rubus smithii Backer, Schoolfl. Java (191 1)
456. — Rubus sundaicus aucL non Blume:
Backer & Bakh. f., Fl. Java 1 (1964) 516, p.p.
— Type: Backer 33873, Java, lecto.
Rubus maximus Kuntze, Meth. Speciesbeschr.
(1879) 62, 76, nom. illeg., non Marsson (1869).
— Rubus moluccanus var. maximus Kuntze,
Rev. Gen. PI. (1891) 222. — Type: not iden-
tified.
Climbing shrubs. Stems up to 10 m long,
hairy, prickles few to many, 1-2 mm long, red-
dish. Leaves ovate, 7-15 by 4.5-9 cm, not or
shallowly lobed, base subtruncate to cordate, mar-
gin grossly serrate, apex acute to subacuminate,
sometimes obtuse, rather stiffly herbaceous, ner-
vation pedate with 6-10 pairs of nerves, venation
widely transverse, upper surface only hairy on
main nerves, lower surface with scattered semi-
appressed to patent hairs on nerves and veins, two-
coloured. Petiole 1-4 cm long. Stipules rather
persistent, 5-9 by 3-5 mm, pinnatisect to -par-
tite with 3-5 pairs of lobes, those 2-5 by 0.2-
0.5 mm. Inflorescence an overhanging or pendant
compound raceme, 15-35 cm long, with up to
12(— 15) laterals of up to 9 cm, the laterals bearing
up to 7 dichasia or cymes, each with up to 5 flow-
ers. Bracts stipule-like to dentate, up to 6 mm long.
Pedicels 6-10 mm long, densely woolly as are the
other axes in the inflorescence. Flowers function-
ally unisexual, plants ?dioecious, flower buds
broadly ovoid. Hypanthium cup-shaped, 4-5 mm
across, densely woolly outside and also with ap-
pressed, straight hairs. Sepals (broadly) ovate,
3.5-5 by 3-5 mm, apiculate just under the ob-
tuse apex outside, margins entire, indumentum out-
side as hypanthium. Petals obovate, 7-9 by 5-6
mm, apex rounded or emarginate, white. Stamens
70-140, glabrous, filaments up to 3 mm, anthers
1.5-2 mm long, staminodes in female flowers 60-
100, up to 1.5 mm long, with minute anther rudi-
ment. Pistils 30-100, ovaries densely long-hairy
on the backside or glabrous, on slightly elevated,
long-hairy torus, styles 1-3 mm long, pistillodes
in supposedly male flowers as pistils but smaller.
Collective fruits globose, up to 1 cm, compact,
sepals spreading under ripe fruits. Fruits up to 3
mm long when dry, exocarp light red (to black-
ish?), still hairy to glabrous, mesocarp rather thick
fleshy or juicy, endocarp with broad dorsal side.
Distribution - Sumatra, W Java.
Habitat - Forest and forest edges, from sea-level
to 2000 m altitude.
Note - Female flowers have obviously non-
functional, small stamens but in flowers with per-
fect stamens the ovaries are not always reduced in
size. Herbarium study is, therefore, not conclusive
on sex distribution in this species.
43. Rubus sorsogonensis Elmer, Leafl. Phi-
lipp. Bot. 10 (1939) 3777. — Type: Elmer
14607, Luzon.
Climbing shrubs. Stems slightly hairy, prick-
les few, short. Leaves ovate, 11-12 by 6-7 cm,
not lobed, base cordate, margin grossly serrate,
apex acute, herbaceous, nervation pedate with 6-7
pairs of nerves, venation widely parallel, sparsely
hairy above and below. Petiole 1—1.5 cm long.
Stipules lanceolate, up to 11 by 3.5 mm, margins
with one minute tooth, with short hairs and short
glandular hairs outside. Inflorescence a terminal
panicle, many-flowered, branches up to 12 cm, the
last branchings di- or monochasial. Bracts up to 15
by 1.5 mm, slightly lobed. Pedicels 1-1.5 cm
long, sometimes with stalked glands between the
hairs, like in the other axes of the inflorescence.
Flowers bisexual, flower buds ovoid, pointed, no
glands on the flowers. Hypanthium saucer-shaped,
4 mm across. Sepals triangular, 10-12 by 3-4
mm, long pointed, outer ones with a minute tooth
on each uncovered margin. Petals none. Stamens
c. 100, glabrous, filaments up to 8 mm, anthers
0.8 mm long. Pistils c. 30, ovaries glabrous, styles
c. 6 mm long. Fruits not seen.
Distribution - Only known from the type, col-
lected in the Philippines (Luzon, on Mt Bulusan),
at 450 m altitude.
Note - Related to Rubus benguetensis.
44. Rubus sundaicus Blume, Bijdr. (1826)
1111; Miq., Fl. Ind. Bat. I, 1 (1855) 383. —
Type: Reinwardt s.n., Tidore (see note).
Stems appressed-cobwebby, prickles very few,
very small. Leaves ovate, 13-15 by 9.5-11 cm,
shallowly 3-lobed, base cordate, margin rather
coarsely serrate-crenate, apex gradually acuminate,
herbaceous, nervation pedate with 9-10 pairs of
nerves, venation widely transverse, upper surface
only with hairs on midrib, lower surface with rem-
nants of cobwebby felt on nerves and veins. Petiole
4-6 cm long. Stipules not seen, scar distinct.
Inflorescence a terminal thyrse, c. 12 cm long,
with 6-7 dichasial to monochasial laterals, up to
2.5 cm long, partly in the axils of leaves. Bracts
incised from apex. Pedicels up to 1 cm long, hairy
as are all other axes. Flowers bisexual, flower buds
ovoid, pointed. Hypanthium cup-shaped, 5-6 mm
across, densely woolly and with semi-appressed
Kalkman — Rosaceae
283
longer hairs outside. Sepals triangular, 6.5-8 by
3.5-5 mm, long-pointed, outer ones with 2 short
teeth on each uncovered margin, inner ones en-
tire and narrower, indumentum outside as hypan-
thium. Petals 6-7 by 5 mm. Stamens c. 65, fila-
ments up to 4.5 mm, glabrous, anthers 0.5-0.8
mm long, glabrous or with 1-2 long hairs on top.
Pistils c. 30, ovaries with long hairs on the dorsal
side, on hairy torus, styles c. 3 mm long. Fruits
not seen.
Distribution - Only known from two sheets bear-
ing different localities, resp. Tidore (Moluccas) and
Java, but probably originating from one collection
made in the Moluccas.
Note - Resembling Rubus cumingii with which
it may be related.
Subgenus Chamaebatus
Rubus subg. Chamaebatus (Focke) Focke, Bibl. Bot. 72 (1910) 17; Zandee & Kalkman,
Blumea 27 (1981) 75. — Rubus sect. Chamaebatus Focke, Abh. Naturw. Ver. Bremen
4 (1874) 145, 146.
Herbaceous to slightly woody, creeping plants. Leaves simple, reniform to cordate,
not or slightly lobed, rarely more deeply incised, nervation pedate with usually 2 or 3 main
side nerves at the very base of the midrib, each with 2 or more basiscopic laterals, nerves
terminating in the margin. Stipules free, on the junction of stem and petiole, persistent.
Flowers bisexual, solitary, terminal, more rarely 2 or 3 on erect laterals. Hypanthium
saucer-shaped, with prickles outside. Inner sepals narrower than outer ones, uncovered
margins pinnately lobed. Fruits loosely cohering, becoming loose from the elevated torus
when ripe.
Distribution — Few species, with a disjunct area: America (Pacific NW America and
Mexico) and Asia (India, China, Taiwan, Japan, Philippines, Java). Two species in Ma-
le sia.
Note — A derived group of small herbaceous plants with reduced inflorescences.
45. Rubus calycinus [Wall, in litL] ex D. Don,
Prod. Fl. Nepal. (1825) 235; Backer & Bakh. f.,
Fl. Java 1 (1964) 515; Steenis, Mount. Fl. Java
(1972) pi. 46-1. —Dalibarda calycina (D. Don)
Seringe in DC., Prod. 2 (1825) 568. — Type:
Wallich, Gosaingsthan.
Rubus boschianus Zoll., Nat. Tijd.. Ned. Indie 14
(1857) 175, 176. — Rubus calycinus D.Don
forma javanicus Kuntze, Meth. Speciesbeschr.
(1879) 106, nom. nud. — Rubus calycinus
D. Don var. suffruticosus Focke, Bibl. Bot. 72
(1910) 21. — Type: Zollinger 2964, Java.
Main stems creeping, up to 3 m long, rooting
on the nodes, with erect, little or not branched
laterals in the axils of leaves, all stems sparsely
hairy, prickles rather few, small. Leaves reniform,
3-6.5 by 3.5-7 cm, usually shallowly 3-5(-7)-
lobed, base deeply cordate, margin serrate, apex
rounded, rather stiff and brittle when dry, nervation
pedate with 2-3(-4) pairs of main side-nerves,
venation reticulate, upper surface with short patent
hairs on the nerves and with small prickles between
the veins, lower surface with long patent hairs and
with needle-shaped, 2.5 mm long prickles on
nerves and veins below. Petiole 2.5-9 cm long.
Stipules ovate, 9-15 by 6-12 mm, base cordate
to rounded, margin finely fimbriate. Flowers soli-
Fig. 7. Rubus calycinus D. Don. Ml Arjuno, Java.
Photo C. G.G.J, van Steenis.
284
Flora Malesiana ser. I, Vol. 11 (2) (1993)
tary, terminal on the laterals, up to 2 cm long
stalked, rarely also one flower in the uppermost
(reduced) leaf. Hypanthium 3.5-5 mm across, pa-
tently hairy outside and with many needle-like, up
to 2.5 mm long prickles. Sepals elliptic to ovate,
sometimes cordate, 11-16 by 5-13 mm, in fruit
growing to 20 by 15 mm, exposed margins pin-
nately lobed with up to 14 lobes, indumentum
outside as hypanthium. Petals elliptic to ovate,
10-11 by 5-7 mm, white. Stamens 30-40, gla-
brous, filaments up to 6 mm, anthers 0.8-1 mm
long. Pistils 30-50(-70), ovaries glabrous, on
elevated, glabrous torus, styles up to 5 mm long.
Fruits orange to red, mesocarp fleshy, only a thin
layer when dry, stone 3-4 mm long, endocarp
rugulose. - Fig. 7.
Distribution - Himalaya region from Nepal to
Arunachal Pradesh, NE India, N Burma, S China;
Malesia: only E Java.
Habitat - In continental Asia in forests, at
1200-2600 m altitude, in Java in different forest
types, locally gregarious, at 1900-2800 m altitude.
Note - The differences between plants from the
Continent and from Java are very slight, see Zandee
& Kalkman, I.e. The description above is based
on plants from Java.
46. Rubus pectinellus Maxim., M61. Biol.
Acad. St. P6tersb. 8 (1872) 374 (= Bull. Acad.
St. Petersb. 17); Elmer, Leafl. Philipp. Bot. 2
(1908) 448; Merr., Enum. Philipp. Flow. PI. 2
(1923) 229. — Type: Maximowicz s.n., Japan.
Main stems thin and wiry, creeping, up to 2 m
long, rooting, with up to 20 cm long, erect, little
or not branched laterals, all stems patently soft-
hairy, prickles rather few, up to 3 mm. Leaves
reniform, 2-5.5 by 2-6 cm, shallowly 3(-5)-
lobed, base deeply cordate, margin serrate, apex
rounded, thinly herbaceous, nervation pedate with
(l-)2 pairs of main side-nerves, venation widely
reticulate, upper surface long-hairy between the
veins, lower surface long-hairy and with needle-
like prickles on nerves and veins. Petiole (l-)2-6
(-8) cm long. Stipules up to 8 by 10 mm, deeply
digitately divided into up to 8 lobes. Flowers soli-
tary, terminal on the laterals, rarely also 1-2 flow-
ers in the axils of the upper (reduced) leaves, up to
3 cm stalked above uppermost true leaf. Hypan-
thium 3-4(-6) mm across, with long, soft, pa-
tent hairs and many needle-like, up to 4 mm long
prickles outside. Sepals elliptic, 8-10 by 3-10
mm, growing to 12 by 13 mm in fruit, exposed
margins pinnately 5-10-lobed, indumentum out-
side as hypanthium. Petals elliptic, 10-13 by
6-9 mm, white. Stamens 16-30, glabrous, fila-
ments up to 5 mm, anthers 1-1.2 mm long. Pis-
tils 24-40, ovaries with few long hairs, on ele-
vated, hairy torus, the styles 3-4 mm long.
Fruits orange to red, mesocarp thin, juicy, stone
2.5-3 mm long, endocarp smooth at first, later
rugulose.
Distribution - SE China, Taiwan, Japan; Male-
sia: Philippines (Luzon, Mindoro, Mindanao).
Habitat - In the Philippines in primary and sec-
ondary forest, also in mossy forest, altitude 750-
2750 m. In China, Taiwan, and Japan in woods, at
comparable altitude.
Note - The description is based on Philippine
specimens only.
Subgenus Rubus
Subgenus Eubatus auct.
47. Rubus plica tus Weihe & Nees, Rubi Germ.
(1822) 15, t. 1.
Small shrubs. Stems sparsely hairy, prickles
few. Leaves pedately 5-foliolate or 3-foliolate,
petiole 2-4.5 cm long. Stipules on the base of
the petiole, linear, up to 12 by 1.5 mm, persistent.
Leaflets ovate to elliptic, terminal ones largest,
3-5.5 by 2-4 cm, base rounded to slightly cor-
date, margin serrate, apex acute to acuminate, ner-
vation pinnate with 8-10 pairs of nerves, upper
surface sparsely hairy on and between the nerves,
lower surface denser hairy all over. Inflorescence a
terminal thyrse with up to 5 few-flowered mono-
chasia under the terminal flower, the lower laterals
in leaf-axils. Bracts and bracteoles persistent. Pedi-
cels and other axes hairy. Flowers bisexual. Hypan-
thium c. 4 mm across. Sepals equal, ovate-trian-
gular, c. 5 mm long, acuminate, entire. Petals
orbicular, c. 8 mm, pink. Stamens c. 100, glabrous.
Pistils c. 40, glabrous. Fruits not seen.
Distribution - This is one of the 'microspecies'
of the Rubus fruticosus complex. It was introduced
(probably from the Netherlands) in the 19th century
and planted near the top of Mt Pangerango in West
Java. Specimens have been collected several times,
possibly always from the same bush. It may still
be alive, but the most recent collection seen was
Kalkman
Rosaceae
285
made in 1931. Fruits were never collected and the
plants may be sterile. The description was made
from the Java plants only.
Note - Other microspecies may also have been
introduced from Europe.
INCOMPLETELY KNOWN SPECIES
Three species of which material is not yet satis-
factory, are mentioned by Kalkman, Blumea 29
(1984) 381-382. They belong to subgenus Mala-
chobatus.
DUBIOUS NAMES
Rubus chartaceus Kuntze, R. edanoi Merr., R.
grewiaefolius Koord. ex Focke, R. guttans Focke,
R. koordersii Focke, R. peltinervius Focke, R. rein-
wardtii Kuntze, and R. zambalensis Elmer are du-
bious names for species reported from Malesia.
Types of these species were either not seen or are
insufficient.
Rubus diclinis F. Muell. var. papuanus Focke.
Neotypification with Brass 30932 not accepted
as in serious conflict with protologue. The speci-
men mentioned is Rubus trigonus.
Rubus mindanaensis Focke, Ann. Cons. Jard. BoL
Geneve 20 (1917) 104.
Considered by Focke to be related to R. niveus
which, according to the material seen, is in die Phi-
lippines restricted to Luzon. Type (W'arburg 14483)
not seen.
Tribus Potentilleae
Herbs, rarely shrubs. Leaves pinnate, trifoliolate, or palmate. Stipules adnate to petiole.
Epicalyx present. Pollen opercular. Few to many pistils on a mostly elevated torus, not
entirely enclosed by hypanthium. Ovule 1, pendulous. Achenes. x = 7.
FRAGARIA
Fragaria L., Sp. PI. (1753) 494; Kalkman, Blumea 16 (1968) 349. —Type species: Fra-
garia vesca L.
Herbs, mostly with stolons. Leaves trifoliolate. Inflorescences cymose, few-flowered.
Flowers 5-merous. Petals white. Stamens many. Pistils many, style ventral. Fruits on en-
larged, fleshy torus, forming a spurious fruit.
Distribution — About 8 species, in Northern temperate regions and in Central and
South America. Not indigenous in Malesia.
KEY TO THE SPECIES
la. Flowers small: hypanthium up to 3 mm diam., sepals 3-4 mm long, petals 4-6 mm
long 2. F. vesca
b. Flowers larger: hypanthium 4-6 mm diam., sepals 7-12 mm long, petals 9-12 mm
long 1. F. x ananassa
1. Fragaria vesca L., Sp. PI. (1753) 494; fruits obovoid, up to c. 1.5 by 1 cm, achenes not
Backer & Bakh. f., Fl. Java 1 (1964) 517.
Leaflets 1.5-3 by 1-2.5 cm, terminal petio-
lule — 1(— 2) mm. Hypanthium 2.5-3 mm diam.
Epicalyx leaves and sepals 3-4 mm long. Petals
4-6 mm long. Stamens 20 or less. Spurious Guinea, but possibly also elsewhere.
sunken in fleshy torus.
Distribution - Temperate Eurasia; in Malesia
introduced and cultivated (not commercially), in
some places escaped and naturalized, seen from Su-
matra, Malaya, Java, Philippines, and Papua New
286 Flora Malesiana ser. I, Vol. 11 (2) (1993)
2. Fragaria x ananassa (Duch.) Guides, fruits globose, ovoid, or obovoid, up to 2 by 1.5
Taxon 33 (1984) 724, and see also the editorial cm, achenes sunken in fleshy torus,
note to this paper by D.H. Nicolson; Backer & Distribution - Cultigen, dispersed all over the
Bakh. f., Fl. Java 1 (1964) 517. — Fragaria world, also running wild. In Malesia a female form
chiloensis (L.) P. Miller x Fragaria virginiana not or rarely setting fruit (see Kalkman, Blumea
P. Miller. 16, 1968, 352) naturalized on Mt Pangerango, W
Java, where it was introduced around 1840.
Leaflets 2-7 by 1.5-6 cm, terminal petiolule Uses - Commercially grown for its fruits, e.g.
up to 9 mm. Hypanthium 4-6 mm diam. Epicalyx in Java (Dieng) and Sumatra (Berastagi); see Choo-
leaves 5-8 by 2.5-3.5 mm. Sepals about equal pong Sukumalanandana & E.W.M. Verheij in
to epicalyx or longer, 7-12 by 3-4.5 mm. Petals E.W.M. Verheij & R.E. Coronel (eds.), Plant Res.
9-12 by 9-12 mm. Stamens 25-40. Spurious SE Asia (PROSEA Handbook) 2 (1991) 171-175.
POTENTILLA
Potentilla L., Sp. PI. (1753) 495; Kalkman, Blumea 16 (1968) 325-354; ibid. 34 (1989)
143-160. — Type species: Potentilla reptans L.
Duchesnea J.E. Sm., Trans. Linn. Soc. 10 (1811) 372. — Type species: Duchesnea fra-
giformis J.E. Sm., nom. illeg. = Fragaria indica Andrews = Duchesnea indica Focke
in Engler & Prand.
Herbs of different habit, rarely shrubs. Leaves compound (palmate, pinnate, trifoliolate,
rarely unifoliolate). Stipules adnate to the petiole, herbaceous or membranous. Flowers
solitary and axillary or opposite the leaves, or in cymose or thyrsoid inflorescences, 5-
(4-6)-merous, bisexual. Hypanthium (shallowly) cup-shaped, lined inside by a some-
times hairy disc. Epicalyx leaves often incised. Sepals valvate, usually entire. Petals entire,
yellow or white, rarely red or purple. Stamens many (up to c. 30) to few. Pistils many to
few, on low to elevated torus; ovaries superior, 1-locular; style apical, ventral or basal,
persistent or jointed and deciduous; ovule 1, pendulous or ascending, inserted near the
place of style-insertion. Fruits free, dry achenes or mesocarp slightly fleshy, surrounded
by persistent epicalyx and calyx, torus rarely enlarging and becoming spongy to fleshy.
Seed with thin testa. — Figs. 8, 9.
Distribution — About 250-400 species, worldwide but mainly in the Northern hemi-
sphere. In Malesia 18 species with a centre of diversity (14 species) in New Guinea.
Habitat — Plants from open and sunny, often wet places. In Malesia most species
montane to subalpine or alpine, at altitudes between 2400 and 4600 m, two species (P. in-
dica and P. sundaica) between 800 and 2400 m.
Ecology — The (sub)alpine species in New Guinea often growing in cushions.
Note — Sometimes parts of the genus as delimited here, are regarded as separate genera,
e. g. Argentina (to which several Malesian species probably would have to be referred if
the genus were accepted), Comarum (not in Malesia), and Duchesnea. The latter two dif-
fer from Potentilla in developing a swollen, spongy torus. It seems a certainty to me that
this is a polyphyletically acquired character state. Separation and recognition of the two
genera leads to a paraphyletic genus Potentilla. Also for Fragaria, and for the same rea-
sons, the combination with Potentilla could be considered. This would, however, either
necessitate hundreds of new combinations in Fragaria, or in the case of conservation, new
and unfamiliar names for the stawberries. (See also Kalkman 1968).
Kalkman — Rosaceae 287
KEY TO THE SPECIES
la. Leaves pinnate 2
b. Leaves ternate or palmate 17
2a. Flowering stems as long as or distinctly longer than the leaves. Stamens usually 10 or
more (often 5 in P. parvula) 3
b. Flowering stems shorter than the leaves. Stamens usually in one whorl of 4 to 6 (10-
15 in P. gorokana) 10
3a. Leaflets densely silky hairy below, leaf surface not visible between the hairs . . 4
b. Leaves glabrous to hairy, but not densely silky, leaf surface visible 7
4a. Stems, petioles, and rachis patently hairy. Flowering stems procumbent
7. P. hooglandii
b. Stems, petioles, and rachis appressed hairy 5
5a. Ovaries hairy 2. P. borneensis
b . Ovaries glabrous 6
6a. Epicalyx leaves entire or practically so. Leaflets crowded, stiff 1. P. adinophylla
b. Epicalyx leaves pinnatifid to pinnatisect. Leaflets distant, not stiff 13. P. papuana
7a. Lateral leaflets entire or some with one incision, apical leaflet bifid . 9. P. indivisa
b. Leaflets incised 8
8a. Leaflets pseudodigitate (pinnately incised but with a minimally short midrib
11. P. linilaciniata
b. Leaflets pinnatisect to pinnatipartite 9
9a. Upper leaflets normally shorter than 10 mm, leaf not more than 10 cm long. Stamens
as many as petals or twice that number, rarely more (-20) 14. P. parvula
b. Upper leaflets 10-30 mm, leaf 9-35 cm long. Stamens 4 times the number of pet-
als or more 15. P. polyphylla
10a. Leaflets digitate, incisions going to the very base 6. P. habbemana
b. Leaflets entire, bipartite, or pinnately incised 11
1 la. Leaflets pinnatisect to pinnatipartite 12
b. Leaflets undivided or bipartite to the very base 16
12a. Epicalyx leaves entire or apically shallowly notched 13
b. At least part of the epicalyx leaves pinnatifid to pinnatipartite, or 3-4-partite . . 15
1 3a. Leaflets densely silky hairy below, leaf surface not visible between the hairs . . 14
b. Leaflets with long hairs below but not densely silky, leaf surface visible
4. P. foersteriana
14a. Leaflets 4-5 pairs 12. P. mangenii
b. Leaflets 12-18 pairs 18. P. wilhelminensis
15a. Leaflets 3-6 pairs. Stamens same number as sepals 10. P. irianensis
b. Leaflets 14-18 pairs. Stamens 2 or 3 times the number of sepals 5. P. gorokana
16a. Leaflets 8-13 pairs 16. P.simulans
b. Leaflets 1-5 (-8) pairs 3. P. brassii
17a. Leaves 5-foliolate. Flowers in dichasial, terminal inflorescences 17. P. sundaica
b. Leaves trifoliolate. Rowers solitary, opposite normal leaves on prostrate stems
8. P. indica
288
Flora Malesiana ser. I, Vol. 11 (2) (1993)
1. Potentilla adinophylla Merr. & Perry, J.
Arnold Arbor. 21 (1940) 190; P. van Royen,
Alpine Fl. New Guinea 4 (1983) 2447, f. 719.
— Type: Brass 1308, Mt Albert Edward.
Rosette plants with stout taproot, leaves and in-
florescences erect Leaves pinnate, up to 7 cm long,
petiole 0.5-1 cm. Stipules membranous. Leaflets
12-18 pairs, crowded and often folded along mid-
rib, up to 8 by 6 mm, serrate to pinnatisect with
3-8 pairs of incisions, densely sericeous below,
less densely above; no intermediary leaves. Flow-
ering stems almost scapose, erect, with 1 or 2
flowers, 4-12 cm long, sericeous. Hypanthium
3.5-4.5 mm diam., densely sericeous outside.
Epicalyx leaves elliptic to ovate, 2-3.5 by 1.2-2
mm, entire or apically notched, sometimes with
shallow incisions, densely sericeous outside. Sepals
triangular, 2.5-4 by 1.2-2 mm, indumentum as
epicalyx. Petals not seen. Stamens 10-20, fila-
ments up to c. 1.5 mm, anthers c. 0.5 mm long.
Torus elevated, densely hairy. Pistils many, ovary
glabrous on hairy stalk, style inserted in the mid-
dle. Achenes c. 1.2 by 0.8 mm, brown, smooth.
Distribution - New Guinea.
Habitat - In grassland, 3100-4100 m altitude.
Note - Some ten specimens known from moun-
tains in Papua New Guinea (Central and Milne Bay
Prov.) and two from Mt Ngga Simanggela (Door-
man) in Irian Jaya, the latter with a less rigid habit.
2. Potentilla borneensis (Stapf) Kalkman,
Blumea 16 (1968) 332. — Potentilla leuconota
D.Don var. borneensis Stapf, Trans. Linn. Soc.
II, 4 (1894) 146. — Type: Haviland 1058, Mt
Kinabalu.
Potentilla leuconota auct. non D.Don: Steenis,
Bull. Jard. Bot. Buitenzorg III, 13 (1934) 242;
Merr., Not. Nat. Acad. Nat. Sc. Philad. n. 47
(1940) 3.
Rosette plant, taproot firm. Stems, petioles,
rachis, underside of leaflets, and pedicels densely
sericeous. Leaves pinnate, 5 — 18(— 32) cm long,
petiole l-4(-9) cm. Stipules membranous. Leaf-
lets 7-16(-24) pairs, elliptic to oblong, gradually
larger to the tip, up to 7-19(-21) by 4-9(-10)
mm, pinnatifid to pinnatisect, rarely incised to near
the midrib, with (4— )6— 1 1 incisions on either
side; intermediary leaflets present or not. Flower-
ing stems 1 to several, erect or ascending, (3-)5-
30(-40) cm, cauline leaves few or none. Thyrse
with up to 13 flowers. Hypanthium 2.5-3.5 mm
diam., densely sericeous outside as are epicalyx and
sepals. Epicalyx leaves elliptic or ovate to oblong,
2.5-5 by (0.7-)l-1.7 mm, apically notched or
entire, rarely deeper incised. Sepals triangular to
triangular-ovate, 2.5-4.5 by 2-3.2 mm. Petals
elliptic to suborbicular, 6-8.5 by 4-6.5 mm,
yellow. Stamens 20, filaments up to 2 mm, an-
thers 0.5 to 0.7 mm. Torus low, hairy. Pistils few
to many, ovary hairy on a hairy stalk, style lateral.
Achenes up to c. 2 by 1.5 mm, brown, smooth
with some veins.
Distribution - Sumatra (only seen from Aceh),
Borneo (only seen from Mt Kinabalu).
Habitat - In stony places on and between rocks,
sheltered or open, in heath-like vegetation or shrub-
land, sometimes in swampy places. In Aceh col-
lected at 2500-3500 m altitude, on Mt Kinabalu
at 3500-4000 m.
Notes - Closely related to P. leuconota from
the Himalayas and Taiwan, and to P. papuana from
Celebes and New Guinea. For the distinction as
species, see Kalkman (1968).
Recently Sojak (Preslia 64, 1992, 221) separated
the Sumatran material as P. sumatrana Sojak. Rec-
ognition on a varietal level would have been more
acceptable, as the differences between the popula-
tions from Aceh and Mt Kinabalu are either over-
lapping or of minor importance.
Some Sumatran specimens have deeply incised
leaflets and may represent a variety (see also note
under P. papuana).
3. Potentilla brassii Merr. & Perry, J. Arnold
Arbor. 21 (1940) 185. — Potentilla foersteriana
Laut. var. brassii (Merr. & Perry) Kalkman,
Blumea 16 (1968) 343; P. van Royen, Alpine
Fl. New Guinea 4 (1983) 2435, pi. 173. —
Types: Brass & Meijer Drees 10156, holo; Brass
9427, Brass & Meijer Drees 10390; all Mt Tri-
kora (Wilhelmina).
Potentilla archboldiana Merr. & Perry, J. Arnold
Arbor. 21 (1940) 185. — Types: Brass & Meijer
Drees 10133, holo; 9839; both Mt Trikora.
Small and compact rosette- herbs, growing indi-
vidually or in cushions up to 60 cm diam. Leaves
pinnate, 1-1.5 cm long, petiole very short. Stip-
ules membranous, hairy outside. Leaflets 1— 5(— 8)
pairs, bipartite or entire, 2.5-5 by 0.5-2 mm,
hard and stiff, glabrous to hairy. Flowering stems
less than 1 cm, with 2 reduced leaves under the
terminal flower. Flowers (3-)4-5-merous. Hy-
panthium 1.5-2 mm diam., growing after anthe-
sis, glabrous to sparsely hairy outside. Epicalyx
leaves elliptic, 1.2-1.5 by 0.5-1 mm, entire,
sometimes partly or all bipartite, indumentum as
hypanthium. Sepals triangular, about as long as
Kalkman
Rosaceae
289
epicalyx, acute. Petals (ob)ovate, c. 2 by 1 mm,
yellow. Stamens isomerous with sepals, filaments
very short, anthers c. 0.5 mm. Torus low, hairy.
Pistils 6-15, ovary glabrous on hairy stalk, style
lateral. Achenes 1-1.2 mm long, red.
Distribution - New Guinea, several mountains
in the main range.
Habitat - Only from altitudes above 3350 m.
KEY TO THE VARIETIES
la Leaves up to c. 1 cm long, leaflets 3-5 pairs,
densely hairy below in the middle part
c. var. strigosa
b. Leaves glabrous below or at most with few
hairs 2
2a. Leaflets 2-5(-8) pairs, most bipartite to the
base, rarely partly undivided or with an addi-
tional smaller third lobe ... a. var. brassii
b. Leaflets 1-2 pairs, undivided b. var. simplex
a. var. brassii — Kalkman, Blumea 34 (1989)
150, f. 2. — Potentilla archboldiana Merr. &
Perry.
Often growing in cushions. Leaflets 2-5(-8)
pairs under the apical one, bipartite, usually to the
base, rarely less deeply divided or some of them
undivided, rarely with a third lobe, lobes 2.5-3.5
by 0.5-0.7 mm, glabrous to sparsely hairy. Hypan-
thium c. 2 mm diam., glabrous or sparsely hairy
outside. Petals obovate. Pistils 6-15.
Distribuuon - New Guinea.
Habitat - Usually in boggy places, altitude
3400-4250 m.
b. var. simplex Kalkman, Blumea 34 (1989)
151. — Type: Hope ANU 10832, Mt Jaya.
Cushions. Leaflets 1-2 pairs under the apical
one, glabrous or few hairs on apex and margin,
lateral leaflets undivided, 3-5 by 1-2 mm, apical
leaflet bifid or trifid, up to 6 by 3 mm. Hypanthium
c. 1.5 mm diam., glabrous outside. Petals ovate.
Pistils 9-12.
Distribuuon - New Guinea: Irian Jaya.
Habitat - Vegetation on poor soils, 3350-c.
4000 m altitude.
c. var. strigosa Kalkman, Blumea 34 (1989) 151.
— Type: Mangen 201 1 , Valentijn Mts.
Cushions. Leaflets 3-5 pairs under the apical
one, unequally bipartite to the base, the largest lobe
up to c. 3 by less than 1 mm, glabrous above, den-
sely long appressed hairy below except near mar-
gins, petiole and rachis densely long-hairy. Hypan-
thium c. 2 mm diam., glabrous outside except near
the rim. Petals elliptic-ovate. Pistils 8-10.
Distribution - New Guinea, only seen from
Valentijn Mts, Irian Jaya.
Habitat - Boggy open places, 3340-3500 m.
4. Potentilla foersteriana LauL, Fedde Rep.
13 (1914) 240; Steenis, Bull. Jard. Bot. Buiten-
zorg IE, 13 (1934) 242; Kalkman, Blumea 16
(1968) 341, excl. var. brassii; P. van Royen,
Alpine Fl. New Guinea 4 (1983) 2434, pi. 172,
f. 717, excl. var. brassii. — Type: Keysser 309,
Finisterre Mts.
Rosette plants with firm taproot, solitary or in
cushions. Leaves pinnate, 2-13 cm long, petiole
up to 1.5 cm. Stipules membranous, long-hairy
below, at least in the central part. Leaflets (5-)8-
16 pairs, elliptic to ovate, base rounded to cordate,
often unequal, pinnatisect to pinnatipartite, rarely
with small intermediary leaflets. Flowering stems
short, 1- to 3-flowered. Flowers (4-)5(-6)-merous.
Hypanthium densely long-hairy outside. Epicalyx
leaves elliptic to ovate, 2-4.5 by 1-2.5 mm,
normally undivided, rarely notched apically, usu-
ally with long hairs outside. Sepals triangular, 2-
4.5 by 1-3 mm, acute, indumentum as epicalyx.
Petals elliptic to obovate, 3-7 by 1.5-3.5 mm,
yellow. Stamens 5-10, filaments up to 1.5 mm,
anthers c. 0.8 mm. Torus low, hairy. Pistils 12-
50 or more, ovary glabrous on short hairy stalk,
style lateral. Achenes up to 1.5 mm long, brown
(to black). - Fig. 8.
Distribution - New Guinea.
Habitat - Open vegetation, (2300-)2700-4150
m altitude.
Note - Potentilla brassii was reduced to a vari-
ety of the present species in Kalkman (1968), but
reinstated as a species in 1989.
KEY TO THE VARIETIES
la. Leaflets (5-)8-12 pairs
a. var. foersteriana
b. Leaflets 12-16(-22) pairs . . . . b. var. ima
a. var. foersteriana
(1989) 152, f. 3.
Kalkman, Blumea 34
Rosettes loose to compact, single or in cushions.
Leaves 2-5 cm, with short petiole. Petiole and
rachis almost glabrous to densely hairy. Leaflets
(5-)8-12 pairs, 3.5-7 by 3-6 mm, pinnatisect
290
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 8. Potentilla foersteriana Laut. var. ima Kalkman. Flowering rosette {Kalkman 4642). Photo C.
Kalkman.
to pinnatipartite with (1— )2— 7 incisions on either
side, usually with long hairs at least on margin
and apex, rarely almost glabrous. Flowering stems
very short, at most 3 cm, with only 1 flower, rare-
ly up to 3, peduncle with 1-2 small leaves or
stipular bracts. Hypanthium 3-4 mm diam. Epi-
calyx leaves 2-4.5 by 1-3 mm. Sepals 2-4.5
by 1-3 mm. Petals (3-)4-6 by 1.5-3.5 mm.
Stamens 5. Pistils 12-32.
Distribution - New Guinea, all over the island.
Habitat - Grasslands and other open vegetation,
altitude 3225-4150 m.
b. var. ima Kalkman, Blumea 16 (1968) 342,
pro max. parte; P. van Royen, Alpine Fl. New
Guinea 4 (1983) 2436, pro max. parte; Kalkman,
Blumea 34 (1989) 152, f. 4. — Type: Kalkman
4642, Mt Kerewa.
Solitary growing rosettes. Leaves 3-13 cm,
petiole up to 1.5 cm. Petiole and rachis densely
long-hairy. Leaflets 12-16 pairs, 6-11 by 3.5-8
mm, pinnatipartite with 1-4 incisions on either
side, with long hairs below. Flowering stems short,
up to 5(-7) cm, usually with 2 flowers, peduncle
with 1-2 small leaves. Hypanthium 3-5 mm
diam. Epicalyx leaves 2.5-4 by 1-2.5 mm.
Sepals 2-3 by 1-3 mm. Petals 3-7 by 2-3.5
mm. Stamens 5-10. Pistils many, up to 50 or
more. - Fig. 8.
Distribution - New Guinea, only seen from
several mountains in Papua New Guinea.
Habitat - Often in grassland, altitude (2300-)
2700-3350(-4100) m.
Note - In its original circumscription this vari-
ety also included P. gorokana, now recognized as a
separate species.
Kalkman
Rosaceae
291
5. Potentilla gorokana Kalkman, Blumea 34
(1989) 155. — Type: Hoogland & Pullen 5513,
Mt Kerigomna.
Solitary growing rosettes with stout, woody
taproot, densely silky on petiole, rachis, underside
of stipules, underside of leaves. Leaves pinnate,
5-1 1 cm long, petiole up to 0.5 cm. Stipules mem-
branous. Leaflets 14-18 pairs, elliptic, 7-9 by
4-6 mm, pinnatisect with 2-5 incisions on each
side, rarely with small intermediary leaflets. Flow-
ering stems up to 3 cm, with 1-4 flowers, peduncle
with few small leaves or stipular bracts. Flowers
5- or 6-merous. Hypanthium 3-5 mm diam., den-
sely long-hairy outside. Epicalyx leaves elliptic,
3.5-4.5 by 1.5-3 mm, mostly pinnatifid to pin-
natisect with 1-3 incisions on each side, long-
hairy outside. Sepals triangular, 3-4 by 1.5-3
mm, indumentum as epicalyx. Petals 5-7 by 3-4
mm, yellow. Stamens 10-15, filaments c. 1 mm,
anthers 0.5 mm. Torus low, hairy. Pistils many,
c. 100, ovary glabrous, stalked, style lateral. Ache-
nes c. 1 mm long, brown.
Distribution - New Guinea, the three specimens
seen all from Goroka Subprov., Papua New Guinea.
Habitat - Open places, altitude 2650-3200 m.
6. Potentilla habbemana Merr. & Perry, J.
Arnold Arbor. 21 (1940) 186; P. van Royen,
Alpine Fl. New Guinea 4 (1983) 2452, f. 720.
— Types: Brass 9594, holo; 9553, 9590; all
Mt Trikora (Wilhelmina).
Loose rosettes. Leaves pinnate, (2-)5-8 cm
long, petiole up to 2.5 cm. Stipules membranous,
underside silky in the middle. Petiole and rachis
long silky hairy to glabrous. Leaflets 6-12 pairs,
digitately divided to the base, lobes 3-6, up to
4 by 1 mm, acute to obtuse, entire, lower side sil-
ky in the middle. Flowering stems 1.5-4.5 cm,
1 -flowered, peduncle with 1-2 leaves, silky. Hy-
panthium up to 4 mm diam., outside long-hairy or
only hairs on rim. Epicalyx leaves elliptic to ligu-
late, 2-2.5 by 0.5-1 mm, entire or one apically
notched, glabrous to sparsely hairy. Sepals trian-
gular, 1.5-3 by 1.2-2 mm, indumentum as epi-
calyx. Petals obovate, 3-3.5 by 1.5-2 mm, yel-
low. Stamens 5, filaments 1 mm, anthers 0.5 mm.
Torus low, hairy. Pistils 7-20, ovary glabrous on
hairy stalk, style lateral. Achenes 1.2 mm long,
greyish brown.
Distribution - New Guinea, only seen from Mt
Trikora, Irian Jaya.
Habitat - Boggy places, heath-like vegetation,
3225-4000 m altitude.
7. Potentilla hooglandii Kalkman, Blumea 13
(1968) 339; P. van Royen, Alpine Fl. New
Guinea 4 (1983) 2444, pi. 174. — Type: Hoog-
land & Schodde 7245, Lagaip Valley.
Rosette plants, forming prostrate, not-rooting,
often long and branched runners bearing leaves and
flowering laterals, taproot stout, all stems up to
the pedicels patently long-hairy. Leaves pinnate,
those of the rosette 5.5-21 cm long, petiole up to
4 cm. Stipules membranous, up to 2 cm long, seri-
ceous outside. Petiole and rachis densely covered
with long, soft, patent hairs. Leaflets 1 1-16 pairs,
± elliptic, basal ones 3-6 mm long, larger to the
apex, up to (8-)10-24 by (4— )6— 1 1 mm, pinna-
tifid to pinnatisect with 6-12 incisions on each
side, densely sericeous below, intermediary leaflets
usually present. Flowering stems procumbent, in
axils of rosette and runner leaves, 4-27 cm long,
with a terminal flower and one or few below it,
peduncle with some (reduced) leaves. Flowers 5- or
6-merous. Hypanthium 3-4 mm diam., densely
sericeous outside. Epicalyx leaves elliptic to ovate,
3-6.5 by 1.5-3.5 mm, serrate in upper part,
indumentum outside as on hypanthium. Sepals
(broadly) triangular to ovate, 3-4.5 by 2-3 mm,
usually entire, indumentum as on hypanthium.
Petals (sub)orbicular to obovate, 5-6 by 4-5 mm,
yellow. Stamens 10-12(-c. 20), filaments 1-1.2
mm long, anthers c. 0.5 mm. Torus thin to thick,
hairy. Pistils very many, ovary glabrous, on short
stalk with some hairs, style lateral below the mid-
dle. Achenes 1 by 0.8 mm, brown to dark purplish
with lighter dorsal line, smooth. - Fig. 9.
Distribution - New Guinea, on several moun-
tains.
Habitat - Open grasslands, altitudes 2440 to
3560 m.
8. Potentilla indica (Andr.) Wolf in Asch. &
Graebn., Syn. Mitt.-Eur. Fl. 6, 1 (1904) 661;
Wolf, Bibl. Bot. 71 (1908) 664; Kalkman,
Blumea 16 (1968) 344. — Fragaria indica Andr.,
Bot. Repos. 7 (1807) L 479; Steenis, Bull. Jard.
Bot. Buitenzorg 111,13 (1934) 241; Backer &
Bakh. f., Fl. Java 1 (1964) 517. — Duchesnea
indica Focke in Engler & Prantl, Nat.. Pflanzen-
fam. 3, 3 (1888) 33. — Type: Andrews, I.e.,
L 479, picturing a plant from a garden in Eng-
land, originating from NE Bengal, India.
Fragaria chrysantha Zoll. & Moritzi, Syst. Verz.
(1846) 7. — Duchesnea chrysantha Miq., Fl.
Ind. Bat. I, 1 (1855) 372. — Type: Zollinger
1987, Mt Tangkubanperahu.
292
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 9. Potentilla hooglandii Kalkman. a. Flowering plant, x 0.7; b. flower, x 2; c. flower, halved length-
wise, x 4; d. ovary, x 12; e. fruit, x 12 (a: Hoogland 9693; b-d: Kalkman 4874, e: Kalkman 4745).
Rosette herbs with long, prostrate, partly sym-
podial stems (stolons) bearing leaves and flowers
and also reduced leaves, daughter plants produced
on the nodes bearing reduced leaves. Stems, pedi-
cels and petioles with long, ± patent hairs and usu-
ally many multicellular glandular hairs. Leaves tri-
foliolate, petiole up to 12(— 16) cm. Stipules mem-
branous, c. 1 cm long, long-hairy outside. Leaflets
sessile or shortly petioluled, apical leaflets rhom-
boid to obovate, 1.5-3.5 by 1-2.5 cm, base cu-
neate, apex rounded, serrate in upper part, lateral
leaflets elliptic to ovate, slightly smaller and usu-
ally with unequal base, all leaflets long hairy be-
low and on the nerves also with glandular hairs.
Flowers solitary, seemingly placed opposite the
runner-leaves, rarely 6-merous, pedicels 2-8 cm
long. Hypanthium 2-3.5 mm diam., sparsely
hairy outside. Epicalyx leaves ± obovate, 3-5 by
2-3 mm during an thesis, distinctly growing af-
terwards, with 2-6 incisions in the apical part,
sparsely hairy outside. Sepals narrowly triangular,
4-6 by 2-3 mm during anthesis, indumentum
outside as epicalyx. Petals obovate, 3.5-4.5 by
2-3.5 mm, yellow. Stamens 15-20, filaments
up to 2.5 mm, anthers c. 0.5 mm. Torus elevated,
hairy or glabrous, distinctly enlarging after anthe-
sis. Pistils many, sessile, ovary glabrous, style
inserted laterally above the middle and in anthesis
much longer than the ovary. Collective fruit 5-1 1
mm diam, soft and fleshy, red. Achenes 0.9-1.3
by 0.7-1 mm, red to brownish, smooth or dis-
tinctly rugose to tuberculate.
Distribution - South, Southeast, and East Asia
In Malesia probably indigenous in Java, Lesser
Sunda Islands and Philippines, cultivated as an or-
namental, escaped from cultivation or at least doubt-
fully indigenous in Sumatra, Malaya, and Singa-
pore. Also introduced and established in many
places in Europe, America, and Africa.
Habitat - Mostly in disturbed habitats: road-
sides, plantations, rarely in forest, rather often in
damp or wet places; altitude 700-2300 m.
Uses - Perry & Metzger (Medicinal plants of E
and SE Asia, 1980, 342) report many medicinal
uses from China, especially for bums, bites, boils,
etc. The fruits are almost tasteless but edible; they
are, however, reported to be poisonous when too
many are eaten.
Kalkman — Rosaceae
293
Vernacular name - As a garden plant often
called the Indian strawberry.
Notes - An extensive overview of the variation
was given earlier (Kalkman 1968). Two groups
can be distinguished, one with tuberculate achenes
on a hairy torus, another with smooth achenes on
a glabrous torus, the correlation of achene and
torus characters not being absolute. Plants of the
first-mentioned group have proven to be diploid
2n = 14, plants of the second group were observed
to be dodecaploid 2n = 84. Japanese authors call
them, respectively, Duchesnea chrysaniha and D.
indica. Published chromosome counts were almost
exclusively made from Japanese or cultivated plants
and none from Malesia. Sterile hybrids obviously
occur in the wild in Japan and can also be artifici-
ally made; they are 7x or 8x and have been named
Duchesnea x hara-kurosawae Naruhashi & Sugi-
moto (J. Phytogeogr. & Taxon. 34, 1986, 11-14;
see also Naruhashi et al. in La Kromosomo 11-42,
1986, 1330-1335).
Because of the lack of any karyological evidence
it is not warranted to extrapolate these findings to
areas outside Japan, but chromosome counts of
vouchered, wild growing specimens of both groups
may establish the occurrence of two (sub)species
in Malesia. Plants with tubercled achenes are in
Malesia known from Luzon, Java, Bali, and Timor,
plants with smooth achenes were seen from Luzon
and other islands and seem to be always introduced
or escaped.
For the distinction of Duchesnea as a genus, see
the note on p. 286 under the genus description.
When brought under Potentilla the group with the
tubercled achenes cannot be called P. chrysantha,
that combination being already occupied.
9. Potentilla indivisa Kalkman, Blumea 34
(1989) 155, f. 5. — Type: Mangen 1163, Mt
Trikora (Wilhelmina).
Rosette herbs with stout taproot. Leaves pin-
nate, 9-10 cm long, petiole up to 2.5 cm. Stipules
membranous, densely long-hairy outside. Petiole
and rachis with few long, soft hairs, sticky when
living (?). Leaflets 10-12 pairs, elliptic-ovate,
oblique, 6-8 by 3-4 mm, lateral ones undivided
and entire, rarely bifid with one smaller lobe, api-
cal ones bifid, all leaflets leathery and glabrous ex-
cept few short hairs at apex. Flowering stems up
to 16 cm, branched, with c. 4 reduced leaves and a
number of bracts, with up to 7 flowers, peduncle
and pedicels sparsely hairy. Hypanthium c. 4 mm
diam., hairy outside. Epicalyx leaves elliptic to tri-
angular-elliptic, 2-2.5 by 1-1.5 mm in anthesis.
afterwards distinctly enlarging up to 5 mm, obtuse,
entire, hard, with few hairs outside. Sepals trian-
gular, 4 by 2.5 mm in anthesis, afterwards up to
6 mm long, indumentum as epicalyx. Petals ob-
ovate, 10 by 7 mm, yellow. Stamens 20, fila-
ments c. 1 mm, anthers c. 0.8 mm. Torus hairy.
Pistils 20-25, ovary glabrous on hairy stalk,
style lateral. Achenes c. 1.5 mm long, brown.
Distribution - New Guinea, only one collection
seen from Irian Jaya, N of Mt Trikora.
Habitat - In tussock grassland, 3100 m altitude.
Note - Related to P. parvula, but with entire
lateral leaflets.
10. Potentilla irianensis Kalkman, Blumea 34
(1989) 156. — Type: Hope ANU 16027, Mt
Jaya (Carstensz).
Rosettes growing in cushions of up to 40 cm
diam. and 30 cm high. Leaves pinnate, very small,
0.5-1.5 cm long, petiole up to 2 mm, perpendic-
ular to the stipule. Stipules membranous, sparsely
to densely hairy outside. Petiole and rachis practi-
cally glabrous to long-hairy. Leaflets 3-6 pairs,
elliptic, up to 2-3 by 1-2.5 mm, base rounded,
tripartite to pinnatisect or pinnatipartite with 1-3
incisions on each side, with few hairs, intermedi-
ary leaflets sometimes present. Flowering stems at
most (in fruiting stage) 1.5 cm long, with 1 or 2
flowers and few reduced leaves on the peduncle.
Hypanthium up to 2.5 mm diam., almost glabrous
outside. Epicalyx leaves ovate to obovate, tri- or
quadripartite with incisions from the top, 1.2-1.5
by 0.7-1 mm, with few hairs. Sepals triangular-
ovate, 1-1.5 by 0.7 mm, with some hairs or gla-
brous. Petals elliptic, c. 2 by 1 mm, yellow. Sta-
mens 5(-7), very small. Pistils 8-12, on the hairy
bottom of the hypanthium, ovary glabrous on
hairy stalk, style lateral. Achenes c. 1.2 mm long,
brown, on thick stalks.
Distribution - New Guinea, 6 collections seen,
all from Mt Jaya in Irian Jaya.
Habitat - Alpine grassland and stony places,
3850-4600 m altitude.
11. Potentilla linilaciniata P. van Royen,
Alpine Fl. New Guinea 4 (1983) 2455, f. 721.
— Type: Van Royen 30113.
Solitary rosette herbs with stout, woody tap-
root. Leaves pinnate, 3 — 15(— 30) cm long, petiole
up to 3(-6) cm. Stipules membranous, silky out-
side. Petiole and rachis long appressed-hairy when
young, glabrescent. Leaflets 12-18(-22) pairs,
pseudo-digitately divided with incisions going to
294
Flora Malesiana ser. I, Vol. 11 (2) (1993)
the very short midrib, the 4-9 lobes 3-7 by 0.5-
1 mm, acute, long-appressed-hairy to almost gla-
brous below. Flowering stems 4-20(-35) cm
long, with (l-)2-5(-12) flowers. Flowers (4-)5-
merous. Hypanthium 2-4.5 mm diam., long-hairy
outside. Epicalyx leaves elliptic, 2-4(-5) by
0.5-2 mm, entire or apically notched, rarely
deeper incised, hairy to almost glabrous. Sepals
triangular, 2.5-4(-5.5) by 1-2.5 mm, indumen-
tum as epicalyx. Petals obovate, 5-8.5 by 3.5-5
mm, yellow. Stamens 15-20, filaments up to 1.5
mm long, anthers c. 1 mm. Torus low to elevated,
hairy. Pistils 15-35, ovary glabrous on hairy stalk,
style lateral. Achenes c. 1.2 mm long, lightbrown,
smooth.
Distribution - New Guinea, seen from several
mountains in Papua New Guinea.
Habitat - Grasslands, usually swampy or boggy,
2700-3800 m altitude.
Note - The leaflets are not truly digitate but have
a recognizable, although very short midrib. The
large variation in length of leaf and inflorescence
may hide two varieties (see discussion in Kalkman,
Blumea 34, 1989, 157).
12. Potentilla mangenii Kalkman, Blumea 34
(1989) 158. — Type: Mangen 495bis, Mt Tri-
kora (Wilhelmina).
Cushion-forming rosettes. Leaves pinnate, up
to 1.5-2.5 cm long, petiole less than 0.5 cm.
Stipules membranous, silky hairy outside. Petiole
and rachis long and densely silky hairy. Leaflets
4-5 pairs, elliptic, up to 5-7 by 3-3.5 mm, pin-
natisect with 1— 3(— 5) incisions each side, densely
silky hairy below. Flowering stems up to 1 cm,
1 -flowered, with 2 reduced leaves on the densely
hairy peduncle. Hypanthium 3.5 mm diam., long-
hairy outside. Epicalyx leaves elliptic, 2-2.5 by
c. 1 mm, entire or rarely notched at apex, long-
hairy outside. Sepals narrowly triangular, 2-2.5
by 1.2 mm, indumentum as on epicalyx. Petals
elliptic, 3.5 by 1.5 mm, yellow. Stamens 5, fila-
ments 1 mm, anthers 0.2 mm. Pistils c. 25, on
the hairy bottom of the hypanthium, ovary gla-
brous, stalked, style lateral. Achenes 1.2 mm long,
shining dark purple.
Distribution - New Guinea, only two specimens
seen, both from Mt Trikora, Irian Jaya.
Habitat - Dry, low vegetation, c. 4100 m alti-
tude.
13. Potentilla papuana Focke, Abh. Naturw.
Ver. Bremen 13 (1895) 162; Steenis, Bull. Jard.
Bot. Buitenzorg III, 13 (1934) 243; Merr. &
Perry, J. Arnold Arbor. 21 (1940) 189; P. van
Royen, Alpine Fl. New Guinea 4 (1983) 2449,
pi. 175. — Potentilla leuconota D. Don var.
papuana F. Muell., nomen. — Type: MacGregor
s.n., Owen Stanley Range.
Potentilla leuconota auct. non D. Don: F. Muell.,
Trans. Roy. Soc. Vict. 1, 2 (1889) 5; Steenis,
Bull. Jard. Bot. Buitenzorg HI, 13 (1934) 242.
Rosette herbs, occasionally forming long, pros-
trate stems, taproot stout. Leaves pinnate, (4.5-)
6-15(-20) cm long, petiole up to 5.5(-7) cm.
Stipules of the rosette leaves membranous, light
brown, outside hairy at least in the centre. Petiole
and rachis densely hairy. Leaflets 7-11 pairs, ellip-
tic to oblong or ovate, the upper ones (7— )1 1—22
by (4-)6-13 mm, pinnatifid to pinnatisect with
4-14 incisions on either side, densely silvery seri-
ceous below and less densely above, intermediary
leaves usually present. Flowering stems several
per rosette, prostrate or ascending, (5-) 10- 30
(-45) cm, with some (reduced) leaves with herba-
ceous, green stipules, usually branched with under
the terminal flower 1-6 laterals with 1-3 flowers
each. Bracts leaf-like but small. Flowers 5(-7)-
merous. Hypanthium 4-5 mm diam., densely
sericeous outside. Epicalyx leaves ovate to ob-
ovate, 3-6 by 2-4 mm, growing after anthesis,
pinnatifid to pinnatisect with (1— )3 — 6 incisions
on either side, dnsely sericeous outside. Sepals tri-
angular to ovate, 3-6 by 2-3 mm, indumentum
as epicalyx. Petals elliptic to obovate, 5-8 by
3.5-6 mm, yellow. Stamens 10-30, filaments
1-2 mm, anthers 0.5-0.8 mm long. Torus low
to high, hairy. Pistils very many, ovary glabrous
on a hairy stalk, style lateral at about the middle.
Achenes 1.2-1.5 mm long, brown to dark purp-
lish brown, smooth.
Distribution - Philippines (only one collection
seen, Mt Tabayoc, Luzon), Celebes, New Guinea.
Habitat - Alpine and subalpine grasslands, wet
or dry, more rarely in shrubland, at (2100-)2600-
3900 m altitude.
Notes - Some New Guinean specimens have
deeply incised leaflets, as also occur in Sumatran
specimens of P. borneensis (see note under that
species).
One chromosome count has been made by Borg-
mann (Zs. f. Bot. 52, 1964, 143), 2n = 42. See
under P. parvula.
14. Potentilla parvula Hook. f. ex Stapf in
Hook., Ic. Plant. IV, 3 (Jan. 1894) pi. 2294;
Stapf, Trans. Linn. Soc. Lond. II, 4 (Dec. 1894)
147; Steenis, Bull. Jard. Bot. Buitenzorg III, 13
Kalkman — Rosaceae
295
(1934) 243; P. van Royen, Alpine Fl. New
Guinea 4 (1983) 2441. — Type: Haviland
1057K, Mt Kinabalu.
Potentilla philippinensis Men., Philipp. J. Sc. 29
(1926) 480, Steenis, Bull. JarcL Bot. Buitenzorg
m, 13 (1934) 243. - Type: Clemens 5006, Mt
Pulog.
Potentilla novoguineensis Merr. & Perry, J. Arnold
Arbor. 21 (1940) 187. — Types: Brass 10727,
Lake Habbema, holo; 4229, Mt Albert Edward;
4636, Wharton Range; Brass & Meijer Drees
9863, Mt Trikora.
Lax to compact rosette herbs, rarely with pros-
trate runner-like branches, taproot stout to slender.
Indumentum of vegetative parts variable but usual-
ly sparsely long-hairy. Leaves pinnate, 2-10 cm
long, in shaded places up to 25 cm, petiole 0.3-
2.5 cm long. Stipules of rosette leaves membran-
ous, about halfway adnate. Leaflets 6-21 pairs,
suborbicular to elliptic, ovate, or obovate, up to
3 — 9(-13) by 2-6(-9) mm, pinnatisect to pin-
natipartite with 1— 5(— 8) incisions on either side,
intermediary leaflets mostly present in the larger
plants. Flowering stems several per rosette, 2.5-
20 cm long, in deep shade up to 38 cm, with some
cauline leaves with herbaceous stipules, bearing
up to 4(-6) flowers, usually sparsely long-hairy.
Flowers (4-)5(-6)-merous. Hypanthium 2-4 mm
diam., hairy outside. Epicalyx leaves suborbicular
to elliptic or obovate, 1.5-3.5 by (0.5— )l-3.5
mm, growing after anthesis, entire or notched at
apex, subglabrous to sparsely hairy outside. Sepals
triangular, 1.8-4 by 1-2.5 mm, indumentum as
on epicalyx. Petals elliptic to obovate, up to 8 by
4 mm but usually smaller, yellow. Stamens 5-10
(-20), filaments 1-2 mm, anthers 0.5-0.8 mm
long. Torus cone-shaped, hairy. Pistils many, rare-
ly few, ovary glabrous, on short, ± hairy stalk,
style lateral at about the middle. Achenes up to 1.3
mm long, brown to purplish or blackish, smooth.
Distribution - Borneo (only seen from Mt Kina-
balu), Philippines (seen from some mountains in
Luzon), Celebes (only seen from Mt Rante Mario),
New Guinea.
Habitat - Mostly in alpine or subalpine grass-
lands, wet or dry, more rarely in shaded places like
open forest, forest edges, and shrubland, at (2065 -)
2600- 3800(-4 100) m altitude.
Notes - The habit seems to be related to the light
conditions in the growing place. Solitary growing
plants in open conditions may form compact roset-
tes while in shaded places the rosette is lax and
open, with longer and more slender leaves and in-
florescences. See Kalkman. Blumea 16 (1968) 337.
One chromosome count has been made by Borg-
mann (Zs. f. Bot. 52, 1964, 144, sub P. novogui-
neensis), 2n = 42. The voucher, Borgmann 24, is
not quite homogeneous. See also under P. papuana.
15. Potentilla polyphylla Wall. [Cat (1829)
nr. 1026, nomen] ex Lehm., Nov. et Min. Cogn.
Stirp. Pug. 3 (1831) 13. — Type: Wallich 1026,
Gossain Than.
Potentilla mooniana Wight, Ic. 1 (1840) t. 233,
text p. xlv; Steenis, Trop. Natuur 21 (1932)
101, f. 3, 4; Bull. Jard. Bot. Buitenzorg III, 13
(1934) 243; Backer & Bakh. f., Fl. Java 1 (1964)
518. — Type: Wight, Nuwara Eliya, Sri Lanka.
Rosette herbs with leafy flowering stems. Leaves
pinnate, 9-35 cm long, petiole 1.5-10 cm. Stip-
ules of rosette leaves membranous, hairy outside.
Petiole and rachis hairy. Leaflets 10-20 pairs,
elliptic, up to 10-30 by 5-12 mm, pinnatisect
to serrate with (5-)7-12 incisions on either side,
sparsely long-hairy, intermediary leaflets present.
Flowering stems several per rosette, ascending to
prostrate, 14-45 cm long, with some cauline
leaves with herbaceous stipules, hairy. Inflorescence
a thyrse with a terminal flower and some lateral
monochasia, 4-10 flowers per inflorescence. Flow-
ers 5(-6)-merous. Hypanthium 3.5-5 mm diam.,
sparsely long-hairy outside. Epicalyx leaves ± el-
liptic, 3-4(-5) by 2-4(-5) mm, growing after
anthesis, usually with 1-4 incisions going half-
way or deeper, sparsely long-hairy outside. Sepals
triangular, 3-4. 5(- 5) by 2-4 mm, indumentum
as on epicalyx. Petals obovate to suborbicular,
5.5-6 by 5-6 mm, yellow. Stamens 19-24,
filaments up to 2 mm, anthers 0.5-0.8 mm long.
Torus cone-shaped, hairy. Pistils many, ovary gla-
brous on hairy stalk, style lateral in the middle.
Achenes c. 1.5 mm long, brown, smooth.
KEY TO THE VARIETIES
la Leaflets serrate, the incisions always less than
halfway. Stems, and especially leaf-rachis and
pedicels soft-hairy, the hairs usually distinctly
spreading. Intermediary leaflets often large,
not rarely 2 or 3 between two panmary leaf-
lets b. var. polyphylla
b. Leaflets pinnatisect, incised about halfway.
Leaf-rachis and pedicels appressed soft-hairy,
sometimes rather sparsely so. Intermediary
leaflets present, small, never more than one
between two primary leaflets.
a. var. kinabaluensis
296
Flora Malesiana ser. I, Vol. 11 (2) (1993)
a. var. kinabaluensis (Stapf) Kalkman, Blumea
16 (1968) 339. — Potentilla mooniana Wight
var. kinabaluensis Stapf, Trans. Linn. Soc. Lond.
II, 4 (1894) 146; Steenis, Bull. Jard. Bot. Bui-
tenzorg III, 13 (1934) 243. — Type: Haviland
1056, Mt Kinabalu.
Distribution - Borneo, only seen from Mt Kina-
balu.
Habitat - Sheltered, damp to peaty places, alti-
tude c. 3300-4000 m.
b. var. polyphylla
Distribution - India, Nepal, Sikkim, Pakistan,
Sri Lanka; Malesia: Java (Mt Papandayan).
Habitat - In Java in herbaceous vegetation, un-
der Anaphalis javanica and Alchemilla villosa, alti-
tude c. 2400 m.
Note - This variety has never been re-collected
in Java since 1940, nor has it been collected in Su-
matra as could be expected. One wonders about the
possibility of its having been intentionally planted,
but Mt Papandayan is not known as a place where
plants have been introduced, like Mt Gede.
16. Potentilla simulans Merr. & Perry, J.
Arnold Arbor. 21 (1940) 187; Kalkman, Blumea
16 (1968) 341, in syn. sub P . foersteriana; ibid.
34 (1989) 159. — Type: Brass 9594A, Mt Tri-
kora (Wilhelmina)
Loose rosettes. Leaves pinnate, up to 5 cm long,
petiole up to 8 mm. Stipules membranous, hairy
outside. Petiole and rachis densely long-hairy.
Leaflets 8-13 pairs, bipartite to the very base,
lobes divergent, elliptic, unequal, the largest one
up to 3.5 by 1.5 mm, with long appressed hairs
underneath, especially in the middle. Flowering
stems up to 1.5 cm long, 1 -flowered and with
some small leaves, densely hairy. Hypanthium c.
2 mm diam., long-hairy outside. Epicalyx leaves
elliptic, entire or one shallowly notched, 1.5-2
by 1 mm, long-hairy outside. Sepals triangular,
1.5-2 by 1.5 mm, indumentum as epicalyx.
Petals broadly obovate, 2-3 mm long, yellow.
Stamens 5, filaments 0.5 mm, anthers 0.5 mm.
Pistils 10-12, on the bottom of the hypanthium,
ovary glabrous, on hairy stalk, style lateral.
Achenes c. 1.2 mm long, shining dark purple.
Distribution - New Guinea, only seen from
Irian Jaya.
Habitat - Grassland on peaty soil, altitude 3225-
4350 m.
Note - Two of the three specimens seen were
taken from mixed collections so the epithet seems
to be quite appropriate.
17. Potentilla sundaica (Blume) O. Kuntze,
Rev. Gen. PL 1 (1891) 219; Steenis, Bull. Jard.
Bot. Buitenzorg m, 13 (1934) 243; Backer &
Bakh.f., Fl. Java 1 (1964) 518. — Fragaria
sundaica Blume, Bijdr. (1826) 1106. — Duches-
nea sundaica Miq., Fl. Ind. Bat. I, 1 (1855) 372,
L 6, incl. var. hirsuta Miq., I.e. 373. — Type:
Blume s.n.,Ui Gede, L sheet nr 909.111-40.
Potentilla kleiniana Wight & Am. in Wight, 111.
Ind. Bot. 1 (1831) t. 85. — Type: Wight 914,
India.
Loose rosettes with erect to prostrate, often
rooting, flowering stems, stolons (prostrate stems
without normal leaves) sometimes present. Leaves
palmately (pedately) 5-foliolate, petiole 6-40 cm
long. Stipules 1.5-4 cm long, membranous, lit-
tle hairy outside. Petiole sparsely to densely soft-
hairy. Leaflets obovate, terminal one shortly peti-
oluled, 2-4.5 by 1-2.5 cm, lateral ones smaller,
coarsely serrate, pinninerved, with few soft appres-
sed hairs underneath. Flowering stems up to 45
(-60) cm, with the cauline leaves smaller than the
rosette leaves and with herbaceous stipules, rarely
3-foliolate, stems and pedicels sparsely to densely
soft-hairy. Daughter plants sometimes developing
on the nodes of the stems. Inflorescences terminal
and axillary, compound dichasia, whole inflores-
cence up to 5-10 cm long. Bracts and bracteoles
leaf-like. Hypanthium 2-3 mm diam., sparsely
hairy outside. Epicalyx leaves ± oblong, 2-3 mm
long, entire or bifid, sparsely hairy outside. Sepals
triangular, 2-3 by 1-2 mm, growing after anthe-
sis, indumentum as epicalyx. Petals obovate, 3-
3.5 by 2-3 mm, yellow, falling early. Stamens
20 or few missing, filaments c. 1 mm, anthers c.
0.5 mm long. Torus high and thin in anthesis,
thicker under the fruit, with few hairs. Pistils
many, ovary glabrous on hairy or glabrous stalk,
style terminal to subapical. Achenes 0.7-1 mm
long, redbrown to brown, rugose.
Distribution - India, Sri Lanka, S China, N
Vietnam, Laos, Korea, Japan; Malesia: only in N
Sumatra and W and C Java.
Habitat - Damp or marshy places like banks of
brooks and lakes, also on roadsides and in villages,
in Sumatra and Java at altitudes of 1200-2300 m.
18. Potentilla wilhelminensis P. van Royen,
Alpine Fl. New Guinea 4 (1983) 2440, f. 718.
— Type: Vers tee g 2534, Mt Trikora (Wilhel-
mina).
Kalkman — Rosaceae
297
Potentilla prob. nov. spec. Kalkman, Blumea 16
(1968) 348.
Rosette herbs, taproot stout. Leaves pinnate,
4-8 cm long, petiole up to 0.5 cm. Stipules mem-
branous, silky outside. Petiole and rachis densely
appressed-silky. Leaflets 12-18 pairs, suborbicular
to elliptic, up to 5-8 by 4-5 mm, stiff and fold-
ed, pinnatifid to pinnatisect with 2-4 incisions
on either side, long silky on both sides. Flowering
stems up to 3 cm, with 1-4 flowers and some re-
duced leaves. Hypanthium hairy outside. Epicalyx
leaves elliptic, 2-3 mm long, entire or apically
notched, hairy outside. Sepals triangular, 2-3 mm
long, indumentum as epicalyx. Petals elliptic, 3-
3.5 by 2-2.8 mm, yellow. Stamens 5, filaments
c. 1.5 mm, anthers c. 0.7 mm long. Torus low,
hairy. Pistils many, ovary glabrous, on hairy stalk,
style lateral. Achenes c. 1 mm long, brown.
Distribution - New Guinea, only seen from Irian
Jaya.
Habitat - Unknown, altitude 4000-4600 m.
Note - The species was collected on Mt Trikora
in 1913 and on Mt Jaya in 1936. Remarkably it
has not been re-collected during more recent explo-
rations of these mountains by G.S.Hope, J.-M.
Mangen and J. Raynal.
Tribus Poterieae
Herbaceous, rarely woody, plants with pinnate leaves. Stipules adnate to petiole. Epicalyx
absent Petals sometimes absent. Pistils several or only 1, superior, enclosed in hypanthium.
Ovule 1, pendulous. Achenes, sometimes drupaceous, x = 7.
ACAENA
Acaena L., Mant. PI. alt. (1771) 200; Bitter, Bibl. Bot. 74 (1911) 1-336. — Type species:
Acaena elongata L.
Herbs, usually creeping, rarely suffrutescent. Leaves imparipinnate, rarely subdigitate.
Inflorescences axillary or terminal, spicate or capitate. Flowers usually 4-merous, bisexual.
Hypanthium ± obconoid, narrowed at the throat, usually armed with few to many barbed
spines. Sepals 3-7, usually 4, valvate, free or shortly connate. Petals absent. Stamens
1-8, often 4, episepalous. Pistils usually 1, sometimes 2-4, free; ovary 1-locular; style
terminal, stigma plumose. Achene(s) remaining enclosed in hypanthium. — Fig. 10.
Distribution — About 40 species (or c. 150 when a much narrower species concept is
followed), almost restricted to the Southern Hemisphere, most richly developed in South
America, going north wards through Central America to California (1 species). Also in
Hawaii (1 species), South Africa (2 species), in the Australian region (several species, in
Australia, Tasmania, New Zealand and neighbouring islands, New Guinea), and on the
subantarctic islands. In Malesia one species in New Guinea.
Acaena anserinifolia (J.R. & G. Forster)
Druce, Bot. Soc. Exch. Club Rep. 1916, Suppl.
2 (1917) 602; Steenis, Bull. Jard. Bot. Buiten-
zorg III, 13 (1934) 241; P. van Royen, Alpine
Fl. New Guinea 4 (1983) 2427. — Ancistrum
anserinifolium J.R. & G. Forster, Char. Gen.
PI. (1776) 2, pi. 2, ' anserinaefolium' . — Type:
Forster s.n., no exact locality.
Prostrate herbs, stems long-hairy. Leaves 2-4
(-9) cm long, petiole up to 0.5(-l) cm. Stipules
membranous, up to 1-1.5 cm long, the free tips
leaflet-like, long-hairy. Petiole and rachis with long
appresscd hairs. Leaflets 4-6 pairs, elliptic to
oblong-lanceolate, upper lateral ones largest, 7-11
(-18) by 2— 5(— 10) mm, serrate, with long appres-
sed hairs underneath. Flowering stems procumbent
298
Flora Malesiana ser. I, Vol. 11 (2) (1993)
1cm
1cm
1cm
b -^k^B^^p'Oj'i
Kalkman — Rosaceae
299
or erect. Inflorescence a solitary, terminal, globose
head, 5-8 mm diam. in anthesis, white, with 50-
80 flowers, peduncle 2-9 cm, hairy, in fruiting
stage head up to 2.5 cm diam. incl. the spines on
the hypanthia. Bracts 7-10, membranous, forming
an involucre close under the head, also some be-
tween the flowers. Flowers bisexual, (sub)sessile.
Hypanthium obovoid, quadrangular in cross sec-
tion, 1-2 mm long, with an ascending spine of
1-2 mm on each of the angles (often one or more
missing), densely long- and stiff-hairy. Sepals 4,
obovate to elliptic, up to 1.8 by 1 mm, erect, per-
sistent. Stamens 2 (or 3), filaments up to 2 mm,
anthers up to 0.5 mm long. Pistil 1, ovary spindle-
shaped, glabrous, only the 1 mm long style pro-
truding, stigma fimbriate. Achene enclosed in the
enlarged hypanthium, the latter then obpyramidal,
quadrangular or with 4 narrow wings, up to 3 mm
long, densely hairy, the barbed spines up to 1 cm
long, reddish to purple. - Fig. 10.
Distribution - Australia from Queensland to S
Australia and Tasmania, New Zealand and nearby
island groups; in Malesia: only in New Guinea.
Habitat - In New Guinea in edges of forest and
shrubland, in open places like landslides and river-
banks, more rarely in subalpine grassland, altitude
1900-4100 m.
Ecology - Epizoochorous by means of the barb-
ed spines. Certainly also anthropochorous.
Notes - Acaena anserinifolia is a complex spe-
cies. Bitter, I.e., who certainly cannot be accused
of irresponsible lumping, placed the many forms
(19 varieties and subspecies) he recognized all in
one species. For New Zealand Allan (Flora New
Zeal. 1, 1961, 361) changed several of those vari-
eties etc. into independent species, admitting that
"the close relationship of these groups is undoubt-
ed." He also remarked (I.e.: 363) that "fertile hy-
brids between them are = common, yet in the ab-
sence of hybridism they are true-breeding,' 7 which
seems to indicate that his species are actually eco-
types.
The description given here is based on New
Guinean material only. This material is very homo-
geneous and matches well the Australian speci-
mens seen.
AGRIMONIA
Agrimonia L., Sp. PI. (1753) 448. — Type species: Agrimonia eupatoria L.
Perennial herbs with creeping rhizomes and erect leafy stems, usually hairy and often
glandular. Leaves interruptedly imparipinnate. Stipules large. Inflorescences terminal,
spike-like racemes. Flowers rather small, 5-merous, bisexual. Hypanthium ± turbinate,
apically armed with erect or patent, hooked bristles, throat almost closed by a dome-
shaped disc with a central pore. Sepais persistent, connivent after anthesis. Petals yellow,
rarely white. Stamens 5-numerous. Pistils 2, free; ovaries 1-locular, style terminal. Usu-
ally one achene developing, enclosed in the indurate, 10-ribbed, armed hypanthium, peri-
carp thin. — Fig. 1L
Distribution — About 15 species, in temperate regions and montane in the tropics, on
the Northern Hemisphere, in S America and S Africa, one species in Malesia (Java).
Agrimonia nepalensis D.Don, Prod. Fl.
Nepal. (1825) 229. — Type: Hamilton s.n.,
Nepal.
Agrimonia blumei G.Don, Gen. Hist. Dichl. PI.
2 (1832) 563, excl. specimens from Japan. —
Agrimonia suaveolens aucL non Pursh (1814):
Blume, Bijdr. (1826) 1113, excl. specimens from
Japan. — Agrimonia javanica Jungh., Java 1
(1853) 664, nom. superfl., illeg.; Miq., Fl. Ind.
BaL I, 1 (1855) 370; Stecnis, Bull. Jard. Bot.
Buitenzorg 17.1, 13 (1934) 241. — Type: prob-
ably Reinwardt 1605 (or 1609 ?), Tengger Mis.
Agrimonia eupatoria L., p.p.: Koord., Exk. Fl.
Java 2 (1912) 333; Backer & Bakh. f., Fl. Java
1 (1964) 519; Steenis, Mount. Fl. Java (1972)
pi. 44-1.
Fig. 10. Acaena anserinifolia (J. R. & G. Forster) Druce. a. Flowering branches: b. leaf with stipules;
c. fruit; d. fruiting head (a, b: Hoogland & Schodde 7619; c, d: Hoogland & Schodde 7428).
300
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Erect herbs, sparsely branched, up to 1 m, stems,
rachises and petioles with long and short patent
hairs, glandular. Leaves 8-16 cm long, petiole 1-5
cm. Stipules herbaceous, amplexicaul, 3 by 3 cm,
deeply serrate, hairy and glandular outside. Leaflets
up to 4 pairs, elliptic to oblong, 2-6 by 1-2 cm,
intermediary leaflets 1-3 between primary ones,
(very) small, all leaflets subsessile, deeply serrate,
appressed- hairy and glandular, also patent hairs on
midrib below. Raceme simple or branched at base,
15-30 cm, in lower part the flowers far apart.
Bracts trifid to tripartite, up to 4.5 mm, bracteoles
2. Pedicels up to 2.5 mm. Flowers staying erect
after anthesis. Hypanthium obconoid, 1-2 mm
long, with 10 obtuse ribs going to the base but
most distinct in upper part, appressed-hairy, under
the sepals with many erect spines, those curved
inwards at apex, up to 1.2 mm. Sepals 2-2.5 by
0.8-1 mm, appressed-hairy mainly on the 3 prom-
inent nerves. Petals elliptic to obovate, 2.5-3.5
by 1.5-2 mm, yellow. Stamens c. 10, filaments
up to 2.8 mm, anthers consisting of 2 subglobular
thecae. Ovary 1 mm, style up to 2 mm. Achenes
usually 1 per flower, fruiting hypanthium 2.5-3
by 3-4 mm, distinctly ribbed, spines all erect and
the larger ones about as long as the calyx. - Fig.
11.
Distribution - Continental SE Asia (Nepal, As-
sam, Burma, S China, N Thailand, Laos, Vietnam);
in Malesia only on some mountains in C and E
Java.
Habitat - Montane, open, often grassy habitats,
altitude c. 1200-2200 m.
Notes - The Java specimens of Agrimonia can-
not be separated from the continental A. nepalensis
D. Don: there is not a single difference.
Some authors place A. nepalensis in synonymy
under A. pilosa Ledeb. (e.g. Nakai, Bot. Mag. To-
kyo 47, 1933, 245; Hara & Kurosawa, J. Jap. Bot.
43, 1968, 392; Yu & Li, Acta Phytotax. Sin. 15,
1977, 89; Purohit & Panigrahi, J. Jap. Bot. 58,
1983, 289) without, however, giving an opinion
about the disposition of the plants from Java.
Other specialists keep the two species mentioned
above separate (Juzepchuk, Fl. U.S.S.R. 10,
1941, 415; Vidal, Fl. Camb., Laos & Vietnam 6,
1968, 134; Skalicky, in litt., 1969). On the basis
of differences in stipules, petals, and indumentum,
and in the absence of a full modern monographic
treatment of the genus over its whole area I prefer
to keep the species separate.
To place A. nepalensis and A. blumei within a
then almost all-embracing species A. eupatoria L.,
as practised in some treatments for Java, does not
seem a sensible classification to me. The most
conspicuous difference between series Eupatoriae
Juz. and Pilosae Juz. is that in the latter the 'fruits'
(fruiting hypanthia) remain erect after anthesis.
There is in BO (and to a lesser degree also in L)
a fair amount of old herbarium material from Java,
but the species has seemingly not been re-collected
since 1941.
Fig. 11. Agrimonia nepalensis D. Don. a. Top of
flowering stem; b. bud; c. flower; d. hypanthium
in fruiting stage {Reinwardt 1605).
Kalkman — Rosaceae 301
SANGUISORBA
Sanguisorba L., Sp. PI. (1753) 116. — Type species: Sanguisorba officinalis L.
Perennial herbs or shrubs, unarmed. Leaves often in rosettes, imparipinnate. Flowers in
terminal spikes or heads on usually long and slender peduncles, 4-merous, bi- or unisexual.
Hypanthium urceolate, constricted at apex, persistent. Sepals deciduous. Petals absent.
Stamens 2-50. Pistils 1 or 2, rarely 3, free; ovaries 1-locular; style terminal, with peni-
cillate stigma. Achenes included in hardened hypanthium.
Distribution — About 20 species, North temperate.
Note — Nordborg, Opera Bot. Lund 11 (2) (1966) 1-103, united not only Sangui-
sorba L. and Poterium L. pro max. p., which is often done, but also Poteridium Spach,
Dendropoteriwn Svent, and Marcetella Svent. The description given here is in accordance
with this large circumscription.
Sanguisorba minor Scop., Fl. Cam. ed. 2, 1 ribbed. Sepals imbricate. Stamens 10-30 in male
(1771) 110; Backer & Bakh. f., Fl. Java 1 and bisexual flowers, filaments up to 8 mm. Pis-
(1964) 519. — Type: Nordborg 8040, Spain, tils 2. Achenes glabrous, pericarp bony in upper
neotype, see Nordborg, Opera Bot. Lund 16 part.
(1967) 98. Distribution - W, C and S Europe, N Africa,
W Asia, introduced and naturalized in N America.
Herb, 0.5 m, stems faintly soft-hairy. Leaves According to Backer & Bakhuizen f., I.e., the spe-
5-9 cm long. Leaflets 3 or 4 pairs, 1.5-2.5 cm cies is in the mountains of Java sometimes culti-
long. Heads globose, up to 1.5 cm diam., with up vated as a pot-herb. I have seen only one sheet of
to 15 crowded flowers, peduncle up to 10 cm long. such a cultivated plant. Also in Europe the leaves
Flowers sessile, bisexual (rarely male) in lower are (were?) used as a vegetable, in soups, and in
part, female in upper part of head. Hypanthium 4- salads.
Tribus Alchemilleae
Herbaceous or shrubby plants with simple, but often deeply palmately incised leaves.
Stipules adnate to petiole. Epicalyx present. Pistils few, superior, enclosed in hypan-
thium. Ovule 1, basal. Achene(s) enclosed in hardened hypanthium. x = 8. Only genus:
Alchemilla s.l., sometimes divided into several genera.
ALCHEMILLA
Alchemilla L., Sp. PI. (1753) 123. — Type species: Alchemilla vulgaris L.
Perennial or annual, erect or prostrate herbs or suffrutices. Leaves simple, usually peti-
oled, ± orbicular in outline, palmately to pedately nerved and palmately incised, folded
in bud. Stipules rather large. Flowers sometimes solitary, usually in cymes, corymbs, or
racemes, small, 4-merous (more rarely 5-merous), bisexual. Hypanthium ± urceolate,
throat almost closed by the disc. Epicalyx leaves usually smaller than sepals, rarely 0.
Sepals valvate. Petals absent. Stamens 1, 2, or 4, rarely 5, inserted outside or inside the
disc, epi- or altemisepalous, short, pollen in many species sterile (apogamy). Pistils 1-4
302
Flora Malesiana ser. I, Vol. 11 (2) (1993)
(-10), free; ovaries sessile or shortly stalked, 1-locular, style ventral or subbasal, protrud-
ing through the disc. Achenes 1-4, remaining enclosed in indurate hypanthium, pericarp
bony to membranous. — Fig. 12.
Distribution — Subcosmopolitan, in all continents. Many species synanthropous and
areas not always entirely natural. In Malesia one species (Java).
Notes — The number of species may be about 200-400 but as in other apogamous
groups it depends very much on the species concept used. Over 300 microspecies have
been described in Europe.
For the description given above, a broad delimitation of the genus is used, i.e. includ-
ing Aphanes, Lachemilla, and Zygalchemilla.
Alchemilla villosa Jungh., Nat. Geneesk. Arch.
Ned. Ind. 2 (1845) 46; Java 1 (1853) 596; Miq.,
Fl. Ind. Bat 1, 1 (1855) 369; Backer & Bakh.f.,
Fl. Java 1 (1964) 518; Steenis, Mount. Fl.
Java (1972) pi. 44-2. — Type: Junghuhn s.n.,
Java, lecto in L, sh. 908.195-1315.
Alchemilla dendroidea Zoll. & Mor. in Zoll., NaL
Geneesk. Arch. Ned. Ind. 1 (1844) 484, nomen,
in obs.
Alchemilla vulcanica Zoll. & Mor. in Zoll., NaL
Geneesk. Arch. Ned. Ind. 2 (1845) 587, nomen,
in obs., non Schlechtend. & Cham. (1830).
Perennial herbs, main stems ± decumbent, older
parts covered with remains of stipules and petioles,
lateral stems prostrate to sub-erect, up to 1 m, not
rooting (?), stems and petioles with patent long
hairs. Stipules of the leaves on the main stems
membranous, those of the leaves of the long
shoots herbaceous, up to c. 1 cm long, shortly
adnate, shortly connate at opposite side of stem,
long-hairy outside. Leaves very variable in size,
blades from 6 by 8 mm to 5 by 8 cm, petioles
1-20 cm. Leaf-blades reniform, (5-)7-9-fid, base
deeply incised, serrate, the apical tooth of each
lobe normally shorter than adjacent ones, pedately
nerved, both sides with semi-appressed hairs. In-
florescences axillary, sympodially stretched cin-
cinni, usually simple, 2-15 cm long, up to c. 12
flowers, peduncle up to 5 cm. Bracts 2 with each
flower, herbaceous. Flowers 4-merous. Hypanthium
narrowly infundibuliform, 1-1.5 by 0.8-1 mm,
densely hairy outside. Disc almost closing throat
of hypanthium, intrastaminal, cushion-shaped.
Epicalyx leaves apert, ovate to elliptic, 0.8-1.2
by 0.5-0.8 mm, hairy outside. Sepals ovate-tri-
angular, 1.2-1.5 by 0.8-1 mm, indumentum as
epicalyx. Stamens 4, alternisepalous, filaments
0.5 mm, anthers c. 0.2 mm, falling early. Pistil 1,
ovary glabrous, shortly stalked, style subbasal.
Achene enclosed in endurated, slightly enlarged hy-
panthium, ovoid, c. 1.2 mm long, style persistent,
pericarp bony, smooth.
Distribution - Malesia: Java, from Mt Papan-
dayan eastwards on many mountains.
Habitat - Grassy places, also in light shade
(Casuarina forest), found at altitudes from 2100 to
3300 m.
Note - The most recent subdivision of the
genus as a whole is to be found in Rothmaler's
paper in Fedde Rep. 40 (1936) 208-212. Rothmaler
did not place A. villosa and its relative from India
and Sri Lanka, A. indica, in section Brevicaules to
which the bulk of the Eurasian species belong, but
in section Longicaules which - apart from the two
Asian species - only contains species from Africa
and Madagascar.
1 mm
Fig. 12. Alchemilla villosa Jungh. Flower (Holst-
voogd 295).
Kalkman — Rosaceae 303
TRIBUS roseae
Woody plants with pinnate leaves. Stipules adnate to petiole. Epicalyx absent. Pistils
many, superior, enclosed in hypanthium. Ovules 1 (or 2), pendulous. Achenes enclosed
in fleshy hypanthium. x = 7. Only genus: Rosa.
ROSA
Rosa L., Sp. P1.Q753) 491; Kalkman, Blumea 21 (1973) 281. — Type species: Rosa
centifolia L. See, however, Taxon 41 (1992) 568 where R. cinnamomea L. is pro-
posed as the type species.
Erect, climbing, or prostrate shrubs, nearly always armed with straight or curved prick-
les, often with glands. Leaves imparipinnate, leaflets pinninerved, usually serrate. Stipules
adnate (rarely, not in Malesia: leaves unifoliolate without stipules). Flowers solitary and
terminal or in terminal thyrses or racemes, large and showy, bisexual, nearly always 5-
merous, cultivars often double. Hypanthium usually globular to urceolate, throat almost
closed by a thickened annular disc. Sepals imbricate, often foliaceous and at least the outer
ones often pinnately incised, persistent or caducous. Petals imbricate, different shades of
red, white, or yellow. Stamens many. Pistils many, rarely few; ovaries superior, shortly
stalked or subsessile, free, included in the hypanthium, 1-locular; styles terminal or lat-
eral, free or with their upper parts coherent to connate, protruding through the hole in the
disc; ovule 1, rarely 2, pendulous. Fruits achenes with usually bony pericarp, included
in the accrescent, ± fleshy, coloured hypanthium (hip). Seed with thin testa; endosperm
absent. — Fig. 13.
Distribution — Probably more than 100 species, in temperate to subtropical regions of
the Northern Hemisphere, some in the montane parts of the tropics. Ornamentals with a
long history of hybridization and with innumerable cultivars of untraceable origin. In Ma-
lesia 2 species indigenous in Luzon, Philippines.
Uses — The modem cultivated roses are almost all complex hybrids. Any of these may
be found cultivated also in SE Asia but they never come beyond the local market. See
D.O.Wijnands in E.Westphal & P.C.M.Jansen (eds.), Plant Res. SE Asia (PROSEA
Handbook). A selection (1989) 240-241.
Note — Occasionally cultivated roses may run wild. See Backer & Bakh.f., Fl. Java
1 (1964) 519-520 and Kalkman, Blumea 21 (1973) 281-291. Only the truly wild spe-
cies are treated here.
KEY TO THE SPECIES
la. Apical leaflets up to 4 cm long. Flowers 7-30, rarely more, in a terminal raceme or
thyrse. Styles hairy 1. R. luciae
b. Apical leaflets up to 1.5 cm long. Flowers usually solitary, terminal on leafy laterals,
rarely also 1 or 2 flowers in the upper leaf-axil(s). Styles glabrous
2. R. transmorrisonensis
304
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Kalkman
Rosaceae
305
1. Rosa luciae Franch. & Rochebr. ex Crepin,
Bull. Soc. Roy. Bot. Belg. 10 (1871) 324;
Kalkman, Blumea 21 (1973) 284, f. 2. — Type:
Savatier specimen in herb. Franchet, Japan.
Rosa wichuraiana Crepin, Bull. Soc. Roy. Bot.
Belg. 15 (1876) 204, nomen; ibid. 25 (1887)
189, descr. — Rosa luciae var. wichuraiana
Koidz., J. Coll. Sc. Imp. Univ. Tokyo 34, art.
2 (1913) 232. — Type: not indicated.
Rosa philippinensis Merr., Philipp. J. Sc. 17
(1921) 260, incl. var. depauperata Merr., I.e.;
Steenis, Bull. Jard. Bot. Buitenzorg III, 13 (1934)
243. — Type: Elmer 5794, holo; several para-
types; all Luzon.
Suberect to straggling or ± climbing shrubs, up
to 4 m. Twigs (scarcely) glandular, otherwise most-
ly glabrous, prickles 2 under each leaf, , up to 4 mm
usually no other prickles present. Leaves 4-10
(-12) cm long, free part of petiole up to 1(-1.5) cm
long. Stipules adnate over (3.5— )8— 1 1 mm, wings
0.7-1.5 mm wide, free tips 1-2.5 mm long, mar-
gin dentate and with stalked glands. Leaflets 7-9,
lateral ones elliptic to elliptic-ovate or elliptic-ob-
long, up to 12-32 by 7-20 mm, apical one larger
than upper lateral ones, margin serrate, glabrous or
with few hairs, lateral leaflets subsessile, apical
petiolules up to 1 cm. Flowers 7-30 or more in a
terminal raceme or thyrse, the lower flowers or
partial inflorescences in the axils of small leaves.
Bracts linear or with expanded apex. Pedicels 1-2.5
cm long, glandular, otherwise glabrous or slightly
hairy. Closed flowerbuds globular, mucronate to
abruptly acuminate, flowers strongly fragrant Hy-
panthium ellipsoid to obovoid, 3-4.5 by 2-3.5
mm in anthesis, outside glandular, with few hairs
or glabrous, long-silky inside. Sepals reflexed
during and after anthesis, caducous, ovate to elliptic-
ovate, acuminate, (6.5-)8-ll by 2.5-4 mm includ-
ing acumen of up to 4 mm, 1-2 side-lobes on ex-
posed margins, glandular and sparsely hairy outside.
Petals (broadly) obovate, 11-18 by 9-11 mm, re-
tuse, white. Stamens 100 or more, filaments up to
7 mm, anthers up to 1.5 mm long. Pistils 12-25,
ovary with a plume of long stiff hairs on one side
at apex, styles 5.5-7 mm long, cohering, hairy.
Hips globular, 6-8(-10) mm, smooth, bluish-
black. Achenes angular ovoid, c. 4.5 mm long, with
hairs on one side, pericarp thick, woody. - Fig. 13.
Distribution - Japan, Korea, Ryukyu Is., E
China, Taiwan; in Malesia: Philippines (Luzon:
several places in the Mountain Prov.).
Habitat- Hardly any information available. Ac-
cording to Merrill, Enum. Philipp. Row. PI. 2
(1923) 231, growing in thickets, 1200-1700 m
altitude.
Note - The description given only pertains to
the specimens from Luzon and does not reflect the
existing variation in the entire area.
2. Rosa transmorrisonensis Hayata, Ic. PI.
Formos. 3 (1913) 97; Kalkman, Blumea 21
(1973) 284, f. 1. — Type: Mori s.n., Mt Mor-
rison.
Rosa luzoniensis Merr., Philipp. J. Sc. 17 (1921)
259; Steenis, Bull. Jard. Bot. Buitenzorg III, 13
(1934) 243. — Types: Santos BS 31876, Mearns
4300, McGregor BS 8336, Pauai.
Climbing shrubs. Twigs thin, glabrous, prickles
2 under each leaf, up to 7 mm. Leaves up to 5 cm,
glabrous, free part of petiole 1-4 mm long. Stip-
ules adnate over c. 5 mm, wings c. 1.2 mm wide,
free tips c. 3 mm long, margins fimbriate and with
glandular teeth. Leaflets (5 or) 7, lateral ones ellip-
tic, apical one larger than lateral ones, 8-15 by 4-6
mm, margin finely serrate, lateral petiolules up to
1 mm, apical one up to 4 mm. Flowers solitary,
terminal on short and leafy laterals, rarely also 1 or
2 in the upper \caf-axi\s.Pedicels up to 1 cm long,
glabrous, not glandular. Closed flowerbuds globu-
lar, mucronate. Hypanthium ellipsoid, c. 4 by 2.5
mm, glabrous and without glands outside, hairy
inside. Sepals reflexed in anthesis, ovate to ellip-
tic, acuminate, 6-7 by 2.5-3.2 mm not includ-
ing the 1-3 mm long acumen, glandular and with
1-3 side-lobes on the exposed margins, glabrous
outside except parts covered in bud. Petals obovate,
up to 11 mm long, retuse, white. Stamens c. 100,
filaments up to 5.5 mm, anthers c. 1.2 mm long.
Pistils c. 12, ovary spindle-shaped, with a plume
of long silky hairs near apex, styles c. 5 mm long,
firmly cohering, glabrous. Hips and achenes not
seen.
Distribution - Taiwan; in Malesia: Philippines
(Luzon: Mountain Prov.).
Habitat - In thickets, altitude c. 1200 m and
higher.
Fig. 13. Rosa luciae Cr6pin. a. Flowering branch, x 1; b, c. flower buds, x 3.5; d. flower halved length-
wise, x 3.5; e. infructescence, x 1 (a: Ramos & Edaho BS 37920; b, e: Jacobs 7595; c, d: Conklin & del
Rosario PNH 72379).
306
Flora Malesiana scr. I, Vol. 11 (2) (1993)
TRIBUS Maleae
Woody plants. Leaves simple, rarely pinnate. Stipules on the very base of the petiole, free
or connate. Hypanthium hollowed, entirely or partly connate with pistils, becoming fleshy.
Epicalyx absent. Carpels (l-)2-5, partly or entirely connate with each other. Ovules 2,
rarely 1, ascending. Pome with bony or membranous endocarp, or multipyrenous drupe
with woody endocarps. x = 17. — Figs. 14, 15.
Fig. 14. Flowers of tribus Maleae, representative Malesian species, slightly idealized, a. Photinia integri-
folia Lindley; b. Photinia davidiana (Decne.) Cardot; c. Eriobotrya bengalensis (Roxb.) Hook, f.; d. Rha-
phiolepis philippincnsis (Vidal) Kalkman; e. Micromeles corymbifera (Miq.) Kalkman (a: Meijer 1669;
b: Fuchs & Collenette 21455; c: Krukoff 4086; d: Sulit PNH 12452; e: Meijer 3466).
Kalian an
Rosaceae
307
Note — The tribe as recognized here confirms to the subfamily Maloideae of most clas-
sifications. In that group often two taxa are recognized, e.g. as Sorbeae and Crataegeae.
Of the genera in Malesia only the non-indigenous Cotoneaster and Pyracantha belong to
the latter group. Iketani & Ohashi in a recent paper [J. Jap. Bot. 66 (1991) 319-351]
recorded the anatomical structure of the fruits of 'Sorbeae' and drew conclusions about
evolutionary trends and phylogenetic relationships. The inclusion of Stranvaesia in Pho-
tinia is supported by them, Micromeles is included in Aria, and Pourthiaea is considered
to be a genus distinct from Photinia [see also Iketani & Ohashi, J. Jap. Bot. 66 (1991)
352-355).
3.rU
Fig. 15. Fruits of tribus Maleae, representative Malesian species, a. Photinia integrifolia Lindley; b. Pho-
tinia davidiana (Decne.) Cardot; c. Eriobotrya bengalensis (Roxb.) Hook, f.; d. Rhaphiolepis philippinensis
(Vidal) Kalkman; e. Micromeles corymbifera (Miq.) Kalkman (a- llochreulincr 908: b: Fuchs & Collenettc
21431: c: Thorenaar 101; d: Sulii PNH 7760; e: Rahmat Si Bocea 10727).
308
Flora Malesiana ser. I, Vol. 11 (2) (1993)
COTONEASTER
Cotoneaster Medikus, Philos. Bot. 1 (1789) 154. — Type species: Cotoneaster vulgaris
Lindley (Mespilus cotoneaster L.).
Evergreen or deciduous shrubs or small trees. Twigs not thorny. Carpels 2-4, free
from each other but connate with the hypanthium, ovary semi-inferior, styles free, ovules 2.
Fruits crowned by persistent sepals, containing 2-4 pyrenes.
Distribution — About 50 species [261 species according to Robertson et al., Syst.
Bot. 16 (1991) 391], centred in E Asia, many species cultivated as ornamentals all over
the world.
Cotoneaster lacteus W.W. Smith, Notes Roy.
Bot. Gard. Edinb. 10 (1917) 23 Clactea'). —
Types: Forrest 10419, 11338, 12720, China.
Shrubs or trees up to 7 m. Twigs hairy when
young, glabrate. Leaves elliptic, 5-6 by 2.5-4
cm, base acute, margin entire, apex rounded and
mucronate, coriaceous, with 6-8 pairs of nerves,
not terminating in the margin, when young sparse-
ly hairy above and densely short-woolly below,
when mature sparsely hairy to glabrous. Petiole up
to 8 mm long. Stipules narrowly triangular, c. 6
by 0.7 mm, hairy outside. Inflorescence a termi-
nal, umbel-shaped, compound raceme, the lower
1 to 3 branches in the axils of (reduced) leaves,
c. 2.5 cm high, 5 cm wide, rachises densely hairy
in anthesis and still hairy in fruit, pedicels up to
4 mm long, densely hairy. Hypanthium and sepals
densely hairy outside, upper rim of hypanthium
free from ovary. Petals orbicular, spreading in an-
thesis, white. Stamens c. 20. Pistils 2, in their
basal half connate with the hypanthium, free from
each other and long-hairy on the free top, styles
terminal. Fruits obovoid, c. 6 by 5 mm when dry
(8 by 6 mm when living), when developing the
free rim of hypanthium and sepals closing around
the top, with 2 stones protruding at the top when
ripe, reddish.
Distribution - Three specimens collected by
J. Sterly in the Gembogl Subprovince, Papua New
Guinea, said to be introduced there from Goroka,
also in Chimbu Province. Cultivated in these places
as an ornamental, but also running wild.
Habitat - At altitudes from 1980 to 2650 m.
Ecology - Fruits eaten and propagated by birds
(Sterly 1751).
Notes - In this predominantly apogamous genus
which, moreover, contains a large number of culti-
vars, species delimitation is difficult. I am by no
means certain that the identification of the three
specimens available is correct, but it did not seem
useful to me to pursue the matter further.
J.M.B. Smith, Science in New Guinea 16
(1990) 13-21, reported the presence in 1989 of
Cotoneaster glaucophyllus on Mt Wilhelm, Papua
New Guinea, as planted and growing well. I did
not see a specimen from that locality but the same
species may be involved as in the other places.
ERIOBOTRYA
Eriobotrya Lindley, Trans. Linn. Soc. Lond. 13 (1921) 102; Blume, Bijdr. (1826) 1102;
Hutch., Gen. Flow. PI. 1 (1964) 214; Vidal, Fl. Camb., Laos & Vietnam 6 (1968)
60; Fl. Thailand 2 (1970) 42; Kalkman, Blumea 21 (1973) 430. — Type species:
Eriobotrya japonica (Thunb.) Lindley.
Unarmed trees or shrubs, evergreen. Leaves simple, margin dentate or entire, main
nerves terminating in the margin. Stipules free or intrapetiolarly connate. Inflorescence a
terminal, compound raceme. Flowers bisexual, 5-merous. Hypanthium obconoid, elon-
gated above the ovary. Sepals persistent. Petals clawed, white. Stamens 15-40. Ovary
semi-inferior to inferior, the hairy top of the connate carpels free from the hypanthium,
Kalian an — Rosaceae
309
2-5-celled, styles as many as cells, usually connate at base, ovules 2 per cell. Fruit a
pome, crowned by the persistent sepals, mesocarp fleshy or with many stone-cells, endo-
carps free from each other, membranous, each containing one seed. Seeds large, testa thin
but hard, endosperm absent, embryo with thick cotyledons. — Figs. 14 c, 15c.
Distribution — About 20 species, from Himalayan region to Japan and throughout SE
Asia southwards to Sumatra, Malaya and Borneo. Only one species indigenous in Malesia.
KEY TO THE SPECIES
la. Leaves soon glabrate, practically glabrous when mature, with 7-10 pairs of nerves.
Stipules free. Petiole 1.5-2.5 cm. Ovary semi-inferior, 2(-3)-celled
1. E. bengalensis
b. Leaves tardily glabrsecent, still densely woolly when mature, with 10-22 pairs of
nerves. Stipules intrapetiolarly cohering or connate. Petiole up to 1 cm. Ovary infe-
rior, 5-celled 2. E. japonica
1. Eriobotrya bengalensis (Roxb.) Hook, f.,
FI. Brit. India 2 (1878) 371; Ridley, Fl. Mai.
Penins. 1 (1922) 681; Prance & Whitmore in
Tree Fl. Malaya 2 (1973) 326, f. 2. — Mespi-
lus bengalensis Roxb., Fl. Ind. 2 (1832) 510.
— Type: Wallich 668/2.
Trees up to 27 m, sometimes shrubs, rarely
buttressed, with spreading branches having flat
foliage ('terminalian' branching), bark rough and
lenticellate, white or grey. Twigs densely hairy
when young, rapidly glabrescent. Leaves oblong to
oblong-lanceolate, 6-17 by 2-6.5 cm, base gradu-
ally tapering, margin shallowly serrate, apex acute
to shortly acuminate, coriaceous, with 7-10 pairs
of nerves, each often with one stronger side-nerve,
those and the primary nerves terminating in the
marginal teeth, venation transverse, almost gla-
brous when mature but with a woolly indumentum
on midrib and nerves when young, red when young
and also when old. Petiole 1.5-2.5 cm, dark. Stip-
ules caducous, triangular, up to c. 4 by 1 mm, free,
ciliolate, sometimes large and semi-foliaceous, up
to 14 by 4 mm. Inflorescence a terminal compound
raceme with up to 12 laterals, the lowermost of
those in the axils of (reduced) leaves, upper ones in
axils of bracts, the panicle up to 14 cm long, pe-
duncle very short, lower laterals up to 12 cm, pedi-
cels 2 — 3(— 5) mm long, densely hairy as are all
other axes in the panicle. Flowers fragrant. Hypan-
thium 1.5-2.5 mm high, densely hairy outside.
Sepals triangular, 2-2.5 by 1-1.5 mm, patent to
reflexed during anthesis, densely hairy outside.
Petals ovate to broadly obovate, 4-4.5 by 2-4.5
mm, reflexed in anthesis, with hairs at base inside,
otherwise glabrous, white. Stamens c. 20, filaments
up to 3 mm, glabrous, anthers 0.6-0.8 mm long.
Ovary inferior when young, later semi-inferior,
2(-3)-celled (see Note), long and densely hairy on
top, styles shortly connate at base, 2-2.5 mm long,
hairy in the lower part. Fruits globular, 10-15 by
9-15 mm (in dry state), exocarp more or less
hairy, grey-green when young, reddish when ripe,
mesocarp hard and gritty, endocarps firm -membra-
nous. Seeds 1 or 2 per fruit, with thin papery testa. -
Figs. 14c, 15c.
Distribution - SE Asia from Eastern part of
Himalayas to Vietnam and West Malesia (Sumatra,
Malaya incl. offshore islands, Bangka?, Borneo).
Habitat - Primary forest, often on limestone,
found from sea-level up to 1200 (to over 1500) m
altitude.
Ecology - Often on limestone, see also Chin
See Chung, The limestone flora of Malaya (1973)
430.
Note - One specimen seen had consistently 5
styles, but this is quite exceptional.
2. Eriobotrya japonica (Thunb.) Lindley, Trans.
Linn. Soc. Lond. 13 (1821) 102; Backer & Bakh.
f., Fl. Java 1 (1964) 512. — Mespilus japoni-
cus Thunb., Fl. Japon. (1784) 206. — Type:
Thunberg s.n., Japan.
Small trees. Twigs rather stout, rough. Leaves
more or less crowded at twig ends, oblong to lan-
ceolate, 12-28 by 3.5-8 cm, margin shorUy den-
tate in upper part, coriaceous, with 10-22 pairs of
nerves, woolly above when very young but soon
310
Flora Malesiana ser. I, Vol. 11 (2) (1993)
glabrate, densely woolly and tardily glabrescent be-
low. Petiole 4-10 mm long. Stipules intrapetio-
lar, cohering or connate into a 2-topped scale, up
to 1 cm long. Inflorescence a compound raceme,
15-20 cm long, peduncle short, pedicels very
short Flowers rather large, very hairy. Sepals per-
sistent. Petals long remaining, white. Ovary infe-
rior or almost so, the densely hairy top free from
the hypanthium, 5-celled, styles (practically) free,
hairy at base. Fruits globular to ovoid, up to 8 cm
diam. but usually (much) smaller, exocarp hairy,
yellow to orange, mesocarp juicy, endocarps mem-
branous. Seeds 2 or 3, large, with firm, glabrous,
brown testa.
Distribution - Native in SE China, there and
in Japan cultivated for many years. Now through-
out the tropics and subtropics cultivated as a fruit
tree. In Malesia: cultivated in home gardens, not
commercially, at medium altitudes.
Uses - The juicy fruits (loquat) are eaten raw
and made into jam. See Nguyen Tien Hiep &
E.W.M. Verheij in E.W.M. Verheij & R.E. Coro-
nel (eds.), Edible fruits and nuts, Plant Res. SE
Asia (PROSEA Handbook) 2 (1991) 161-164.
MALUS
Malus Miller, Gard. Diet. (1754). — Type species: M. sylvestris Miller (Pyrus malus L.).
Trees or shrubs, unarmed or with thorns, mostly deciduous. Leaves simple, lobed or
toothed. Flowers in few-flowered, simple racemes. Hypanthium with a free rim above the
ovary, the rim persistent or rupturing after anthesis. Ovary inferior, carpels completely ad-
nate with the hypanthium and without exposed free top, 3 -5-celled, styles connate at base.
Fruit a pome, mesocarp fleshy, in most species without stone cells, endocarp cartilaginous.
Distribution — About 50 species in Eurasia and N America. In Malesia only a culti-
vated species.
Malus domestica Borkh., Hand. Forstbot. 2
(1803) 1272. — Malus sylvestris Miller. —
Malus pumila Miller.
Small trees, up to 10 m. Twigs hairy when
young. Leaves elliptic-ovate, 4-13 by 3-7 cm,
base rounded, margins serrate, apex acute, with 3-6
pairs of nerves, usually hairy underneath. Inflores-
cence a simple raceme, terminal on short shoots.
Hypanthium hairy outside. Sepals persistent on
the fruit. Petals white to pinkish. Ovary 4- or
5-celled, inferior, ovules 2 per cell. Fruit a pome,
globular to obovoid, exocarp glabrous, variously
coloured, mesocarp fleshy, without stone cells, en-
docarp leathery to bony. Seeds usually 2 per cell.
Distribution - Originated in West Asia, now
spread over the entire world. In Malesia: cultivated
in E Java, Timor, the Philippines, and probably
some other islands.
Uses - See Surachmat Kusumo & E.W.M.
Verheij in E.W.M. Verheij & R.E. Coronel (eds.),
Edible fruits and nuts, Plant Res. SE Asia O^ROSEA
Handbook) 2 (1991) 200-203 for a review of the
cultivation of the apple in Malesian countries.
Note - The name accepted pertains to the culti-
vated apple which is supposed to be derived from
wild Malus pumila and several other species hybrid-
ized with it. Since the genetic make-up of its
many cultivars cannot be ascertained, a separate
specific epithet seems warranted, although other
ways of reasoning may lead to the acceptance of
one of the other names mentioned above.
MICROMELES
Micromeles Decne., Nouv. Arch. Mus. Paris 10 (1874) 168; Kalkman, Blumea 21 (1973)
437. _ py rus l. sect. Micromeles Hook, f., Fl. Brit. India 2 (1 878) 377. — Sorbus L.
sect. Micromeles Rehder, Manual Cult. Trees & Shrubs (1927) 382; Vidal, Fl. Camb.,
Laos & Vietnam 6 (1968) 24. — Type species: not designated.
Aria (Pers.) Host, Fl. Austral. 2 (1831) 7, p.p.: Robertson et al., Syst. Bot. 16 (1991) 389.
Kalkman
Rosaceae
311
Unarmed trees or shrubs, deciduous. Leaves simple, with the main nerves terminating
in the serrate margin. Stipules free. Inflorescence a terminal, panicle-shaped, compound
raceme. Flowers bisexual, 5-merous. Hypanthium obconoid, elongated above the ovary,
the free part transversely rupturing after anthesis and falling with sepals and other flower-
parts, its inside covered with a disc. Stamens c. 20. Ovary inferior, apex covered by the
hypanthial disc, 2-5-, most often 3-celled, styles as many as cells, usually connate at
base, ovules 2 per cell. Fruit a pome, exocarp usually lenticellate, mesocarp hard and dry,
endocarp thin. Seeds several, testa rather thin, endosperm absent, embryo with flat cotyle-
dons. — Figs. 14 e, 15 e, 16.
Distribution — Less than 15 species, in SE and E Asia, one species also in W Malesia.
Note — There is considerable disagreement about the status of this genus: must it be
combined with Pyrus, with Sorbus, with Aria, or be kept separate? For the time being I
prefer to cling to my earlier (1973) decision to keep the genus separate.
Micromeles corymbifera (Miq.) Kalkman,
Blumea 21 (1973) 437. — Vaccinium? corymbi-
ferum Miq., Fl. Ind. Bat. Suppl. 1 (1861) 588.
— Sorbus corymbifera (Miq.) Hiep & Yakovlev,
Bot. J. 66 (1981) 1 188. — Type: Junghuhn s.n.
(PI. Jungh. Ined. 1035), Sumatra.
Pyrus granulosa Bertoloni, Mem. Accad. Sc. Bolog-
na II, 4 (1864) 312, pi. 3; Ridley, Fl. Mai.
Penins. 1 (1922) 680; Steenis, Bull. Jard. Bot.
Buitenzorg III, 13 (1934) 242; Blumea 12
(1964) 14 (transfer of Vaccinium corymbiferum).
— Type: Hooker f. & Thomson 'Pyrus c , India.
Micromeles malayensis Ridley, J. Bot. 62 (1924)
296. — Type: Nur 11241, Malaya.
Photinia bartletlii Merr., Pap. Mich. Acad. Sc. 19
(1934) 155; Steenis, Bull. Jard. Bot. Buitenzorg
III, 13 (1934) 242. — Type: Bartlett 8662, Su-
matra.
Trees up to 30 m, or shrubs, sometimes hemi-
epiphytic, deciduous, bark brown, scaly. Twigs
glabrous or with very few hairs, with light-col-
oured lenticels. Leaves oblong to elliptic-oblong,
rarely ovatish, 7-13 by 4-7.5 cm, base acute,
more rarely rounded, margin shallowly serrate with
usually the basal quarter entire, apex acute to acumi-
nate, herbaceous, with 8-1 1 pairs of nerves, vena-
tion transverse, not very prominent, sometimes
with small, cylindrical glandular outgrowths on
midrib above, very young leaves sometimes (see
Note) ferruginous-woolly, but also in that case ma-
ture leaves entirely glabrous, the indumentum dis-
appearing completely. Petiole 1-3 cm long, red as
are midrib and leaf margin. Stipules early falling,
very small and brisUe-like, but sometimes on the
basal leaves of a shoot well -developed and up to 6
by 1 mm. Inflorescence up to 7 cm long, branched
up to the third order with the lower primary branches
axillary to normal leaves, all axes, including the up
to 5 mm long pedicels, glabrous, sometimes with
few hairs, rarely densely hairy (see Note). Flowers
5-merous, fragrant. Hypanthium obconoid, often
more or less abruptly widened in upper part, 2-3
mm high, the upper 1 mm free from the ovary and
falling after anthesis, glabrous outside or with some
hairs, inside covered by a glabrous disc. Sepals tri-
angular, 1.5-2.5 by 1.5-3 mm, glabrous. Petals
(broadly) elliptic, ovate or obovate, 4.5-6 by 3-
4.5 mm, white, glabrous. Stamens 19-24, fila-
ments up to 6 mm, anthers 0.7-1 mm long. Ovary
(2-)3(-4)-celled, styles distinctly connate in
lower part, up to 5.5 mm long. Fruits globular to
ellipsoid, sometimes more ovoid or obovoid, rare-
lv rjear-shaped (see Note), with a large circular scar
■v7'
Fig. 16. Micromeles corymbifera (Miq.) Kalkman.
Fruiting branch. Mt Sago, Sumatra. Photo W.
Meijer.
312 Flora Malesiana ser. I, Vol. 11 (2) (1993)
at apex, usually still present when young flush ap- ing the tree-habit. It does not seem to be a strangler,
pears, 9-19 by 8-17 mm, exocarp brownish, al- killing its host, as some Ficus do. See Van Steenis,
ways with corky lenticels, mesocarp very hard and Flora Malesiana 1, 4 (1948) xxix, and Corner, Way-
woody. Seeds 1 or 2 per cell, flat, up to 5 by 3.5 side Trees, ed. 3 (1988) 620, sub Pyrus corymbifera
mm, testa firm-membranous, glabrous. - Figs. (nom. illeg., non Nakai, 1935).
14 e, 15 e, 16.
Distribution - Continental Asia: India (Assam), Notes - Some collections have (traces of) a
China (Yunnan), Thailand, Laos, Cambodia, Viet- dense woolly indumentum on inflorescences and
nam; Malesia: Sumatra, Peninsular Malaysia. leaves, but mosUy the specimens are (almost) gla-
Habitat - Primary montane forest, also in mos- brous, the indumentum that may have been present
sy forest, and in more open, shrubby vegetation, at at a young stage having disappeared rapidly and
(600?-) 1 100-3000 m altitude. thoroughly.
Ecology - One of the few examples of the hemi- In only one collection from Sumatra {van Steenis
epiphytic life-style. The plant may start as an epi- 10031) the fruits are pyriform as in var. turbinata
phytic shrub, later sending down roots and acquir- Cardot which is known from the continent.
PHOTINIA
Photinia Lindley, Trans. Linn. Soc. Lond. 13 (1821) 103; Kalkman, Blumea 21 (1973)
418. — Type species: Photinia serrulata Lindley.
Stranvaesia Lindley in Edw., Bot. Reg. 23 (1837) t. 1956. — Type species: Stranvaesia
glaucescens Lindley.
Unarmed trees or shrubs, evergreen or deciduous, Malesian species all evergreen.
Leaves simple, entire to serrate, the secondary nerves not running to the margin. Stipules
free. Inflorescence a terminal, panicle- or corymb-shaped compound raceme. Flowers bi-
sexual, 5-merous. Hypanthium obconoid to campanulate, elongate above the ovary. Sepals
persistent. Petals more or less distinctly clawed. Stamens 16-25. Ovary semi -inferior,
usually hairy on the exposed, free top, (l-)2-5-celled, styles as many as cells, connate at
base or free, ovules 2 per cell. Fruit a pome, covered at apex by the persistent free part of
the hypanthium and the sepals, core consisting of the bony endocarp. Seeds 1 or 2 per
cell, rather small, testa rather hard, endosperm thin or absent. — Figs. 14 a, b, 15 a, b, 17.
Distribution - About 50 species in E Asia, 5 species extending into Malesia.
Note - Robertson et al., Syst. Bot. 16 (1991) 391, included the N American genus
Aronia in Photinia.
KEY TO THE SPECIES
la. Leaves, when dry, with many black to brown glandular dots scattered on the under-
side 4. P. prunifolia
b. Leaves without dark dots 2
2a. Leaves entire 3
b. Leaves crenate to serrate, at least in upper part 4
3a. Ovary 2- (or 3-)celled 2. P. integrifolia
b. Ovary 4- or 5-celled 1. P. davidiana
4a. Inflorescences glabrous. Ovary 2- or 3-celled 5. P. serratifolia
b. Inflorescences rather densely shortly woolly. Ovary 4- or 5-celled ... 3. P. nussia
Kalkman
Rosaceae
313
1. Photinia davidiana (Decne.) Cardot, Bull.
Mus. Nat. Hist. Nat. Paris 25 (1919) 399. —
Stranvaesia davidiana Decne, Nouv. Arch. Mus.
Paris 10 (1874) 179. — Type: David s.n.,
Tibet.
Stranvaesia integrifolia Stapf, Hook. Ic. PI. 23
(1894) L 2295; Steenis, Bull. Jard. BoL Buiten-
zorg m, 13 (1934) 246. — Type: Haviland 1071
K, Mt Kinabalu.
Shrubs or trees, up to 4.5 m. Twigs shortly hairy
when young. Leaves elliptic to elliptic-oblong or
slightly ovate or obovate, 4-9 by 2-4 cm, base
rounded to acute, margin entire, apex usually acute
and mucronate, sometimes shortly acuminate, stiff-
coriaceous, with 7-9 pairs of nerves, often not dis-
tinctly different from strong intermediate veins, ve-
nation widely reticulate, reddish when young, red
before falling, shortly hairy on main nerves when
young, glabrescent. Petiole 0.5-1 cm long. Stip-
ules narrowly triangular, up to 3 by 0.5 mm, keeled
inside, sparsely hairy, glandular on the keel, cadu-
cous. Racemes pyramidal, up to 4 cm long, with
up to 8 first order laterals of up to 3.5 cm long and
usually branched again, lowermost laterals usually
in axils of leaves, rachises hairy, pedicels up to 3
mm long, hairy. Hypanthium campanulate, up to
2 mm high, only the basal half connate with the
ovary, sparsely hairy outside. Sepals triangular, 1-
1.5 by 1.7-2 mm, usually ciliolate. Petals elliptic
to broadly orbicular, up to 4.5 by 4 mm, white to
pinkish. Stamens 17-20, filaments up to 3 mm,
anthers up to 1 mm long. Ovary semi-inferior,
shortly hairy on the dome-shaped free top, 3-?, 4-
or 5-celled, styles up to 3.5 mm long, halfway
connate, hairy at extreme base. Fruits (sub)globu-
lar, up to 8 by 8 mm when dry, free part of hypan-
thium and sepals closely appressed against the top,
exocarp sparsely hairy, red, mesocarp thick and
fleshy, up to 2 mm thick when dry, endocarp thin,
bony. Seeds 1-5, ellipsoid to obovoid, c. 4 by 2
mm, with firm, brown testa. - Figs. 14b, 15b.
Distribution - China, Taiwan, N Vietnam; Ma-
lesia: Sumatra (only few seen, from Aceh), Borneo
(only seen from Mt Kinabalu, rather many collec-
tions).
Habitat - In Malesia in subalpine shrubland, at
2600-3900 m altitude, on the continent lower.
2. Photinia integrifolia Lindley, Trans. Linn.
Soc. Lond. 13 (1821) 103; Blume, Bijdr. (1826)
1103; Miq., Fl. Ind. Bat. I, 1 (1855) 387; Backer
& Bakh. f., Fl. Java 1 (1964) 513. — Type:
Wqllich 669, Nepal.
Photinia notoniana Wight & Am., Prod. 1 (1834)
302; Koord. & Valeton, Bijdr. Booms. Java 5
(1900) 361; Koord., Exk. Fl. Java 2 (1912) 318;
Steenis, Bull. Jard. BoL Buitenzorg III, 13 (1934)
242. — Type: Wight 1014, lecto chosen by
Vidal, from India.
Photinia notoniana Wight & Am. forma grandi-
flora Koord. & Valeton, Bijdr. Booms. Java 5
(1900) 361 — Type: not indicated, described
from Mt Gedeh, Java.
Shrubs or trees up to 15 m. Twigs glabrous or
somewhat hairy when young. Leaves elliptic to
oblong, 4— 15(— 21) by 2.5-8 cm, base cuneate
to rounded, entire (rarely faintly toothed), apex ob-
tuse to acute, usually shortly acuminate, coriace-
ous, with 6-12 pairs of secondary nerves which
are usually not well distinguishable from stronger
tertiary ones, venation widely reticulate, glabrous
or with few hairs when young. Petiole 0.5-4 cm
long, often red as are midrib and leaf mar-
gins. Stipules triangular, up to 2 by 1 mm, early
caducous. Inflorescence up to 7(-12) cm long,
branched up to the 4th order, with up to 12 or
more spreading branches of the first order, the
lower ones often in axils of leaves and up to 8(— 10)
cm long, all rachises up to the 0-7 mm long
pedicels short-hairy or glabrous. Flowers fragrant.
Hypanthium obconoid, 1-2 mm high, usually
glabrous outside. Sepals triangular, obtuse, (0.5 -)
0.8-l(-1.5) by 1-2 mm. Petals (sub)orbicular,
2.5-4 by 2-4 mm, white. Stamens 17-21, fila-
ments up to 2.5(-4) mm, glabrous, anthers 0.5-
0.8 mm long. Ovary usually hairy on the exposed
top, 2- (rarely 3-)celled, styles up to 3 mm, usual-
ly shortly to halfway connate. Fruits subglobular,
4-8 by 3.5-6.5 mm, upper part of hypanthium
and sepals closely appressed against top, exocarp
red, mesocarp fleshy, endocarp hard, bony. Seeds
usually 2 per cell, ovoid, up to 4 mm long, with
hard, brown, glabrous testa, endosperm thin. - Figs.
14 a, 15a, 17.
Note - The species has in various ways been
divided into varieties, see discussion in Kalkman,
Blumea 21 (1973) 418. The two varieties as ac-
cepted here, certainly have no phylogenetic value
whatsoever.
a. var. integrifolia — Kalkman, Blumea 21
(1973)423.
The synonyms given above under the species per-
tain to the type variety.
Twigs always glabrous. Rachises of the inflo-
rescence, including the pedicels glabrous.
314
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 17. Photinia integrifolia Lindley. Fruiting shrub. Mt Arjuno, Java. Photo J. Jeswiet.
Distribution - Continental Asia (Tibet, Nepal,
Sikkim, Bhutan, NE India, S China, Thailand, Sri
Lanka); Malesia: Java, Lesser Sunda Islands.
Habitat - In Malesia in montane forest and sub-
alpine shrubbery, altirude (1400-)2000-3350 m.
Distribution - Continental Asia (NE India, Bur-
ma, S China, Laos, S India, Sri Lanka; Malesia:
Sumatra, Malaya, Java, Lesser Sunda Islands.
Habitat - In Malesia in montane forest and sub-
alpine vegetation, altitude 1300-3200 m.
b. var. sublanceolata Miq., Fl. Ind. Bat. I, 1
(1855) 387; Kalkman, Blumea 21 (1973) 423.
— Type: Horsfield 432, Java.
Photinia dasythyrsa Miq., Fl. Ind. Bat. I, 1 (1855)
387; Steenis, Bull. Jard. Bot. Buitenzorg m, 13
(1934) 242. — Type: fragment in U?
Photinia integrifolia Lindley var. subdenticulata
Miq., Fl. Ind. Bat. I, 1 (1855) 387. — Type:
Horsfield 1135, Java.
Twigs glabrous to distinctly hairy when young.
Rachises of the inflorescence, including the pedi-
cels with a cover of short, appressed hairs which
do not entirely disappear with age.
3. Photinia nussia (D. Don) Kalkman, Blumea
21 (1973) 429. — Pyrus nussia D.Don, Prod.
Fl. Nepal. (1825) 237. — Stranvaesia nussia
(D. Don) Decne., Nouv. Arch. Mus. Paris 10
(1874) 178; Vidal, Adansonia 5 (1965) 231,
577. — Type: Hamilton s.n., lecto, Nepal;
Wallich 658, para.
Eriobotrya ambigua Merr., Philipp. Bur. Gov. Lab.
Pub. 35 (1906) 19; Enum. Philipp. Flow. PI.
2 (1923) 226. — Stranvaesia ambigua Nakai, J.
Arnold Arbor. 5 (1924) 72. — Type: Meyer FB
2796, Luzon; paratypes: Whitford 1155, 1168,
1307, Luzon.
Kalkman
Rosaceae
315
Eriobotrya oblongifolia Merr. & Rolfe, Philipp. J.
Sc, Bot. 3 (1908) 102; Merr., Enum. Philipp.
Flow. PI. 2 (1923) 226. — Type: Mearns &
Hutchinson FB 4680, Mindanao.
Trees up to 10 m. Twigs densely shortly woolly
when young, glabrescenL Leaves oblong to oblong-
lanceolate, 5-11 by 2-4 cm, base acute, margin
entire in lower half, shallowly crenate to serrate in
upper part, apex acute to shortly acuminate, coria-
ceous, with 8-15 pairs of secondary nerves, often
not distinguishable from stronger tertiary nerves,
venation widely reticulate, both surfaces shortly
woolly when young, hairs soon vanishing and
ultimately quite glabrous. Petiole 1-2 cm long.
Stipules narrowly triangular, 1.5 -6 by 0.5-0.8
mm, not very early caducous. Inflorescences cor-
ymb-shaped, 4-6 cm long, with up to 12 primary
branches, the lower ones in axils of leaves and
branched again, up to 6 cm long, all rachises den-
sely short-woolly, including the up to 7 mm long
pedicels. Hypanthium obconoid, 1.5-2.5 mm
high, densely short-woolly outside. Sepals broadly
triangular, 1-1.8 by by 1.5-2.5 mm. Petals ellip-
tic to ovate, 4-5.5 by 3-3.5 mm, white. Stamens
c. 20, filaments up to 3 mm, glabrous, anthers
0.7-1 mm long. Ovary densely hairy on exposed
top, 5- (rarely 4-)celled, styles up to 4.5 mm, half-
way connate, with hairs at base. Fruits globular,
up to 6 mm diam., upper rim of hypanthium and
sepals closely appressed against its bulging, dome-
shaped, now almost glabrous top, the pericarp thin
and brittle when dry, practically glabrous, colour
unknown. Seeds not seen in mature state.
Distribution - Continental Asia (E Himalayas,
NE India, N Burma, N Thailand, N Laos, S China,
but not recorded from Taiwan); Malesia: Philip-
pines.
Habitat - On the continent in evergreen forest
types, altitude 500-1800 m, in the Philippines
hardly any information available, but up to 2500 m
altitude.
Note - The description only pertains to the Phi-
lippine material.
4. Photinia prunifolia ((Hook. & Am.) Lind-
ley in Edw., Bot. Reg. 23 (1837) t. 1956. —
Photinia serrulata (1 prunifolia Hook. & Arn.,
Botany Beechey's Voy. (1833) 185. — Type:
Beechey's Coll. s.n., Macao.
Small trees up to 12 m, bark (dark) brown. Twigs
glabrous. Leaves lanceolate, 9-14 by 2-4.5 cm,
base cuneate, margin irregularly and finely glandu-
lar-serrate except at very base, apex gradually taper-
ing, with 12-16 pairs of secondary nerves, not
always distinguishable from stronger intermediary
tertiary nerves, venation reticulate, glabrous on
both sides, glossy above, with many scattered
brown to black glandular dots on the dull under-
side. Petiole 0.7-1.5 cm long. Stipules subulate,
3-6 mm long, very early caducous. Inflorescences
corymbose in shape, up to 8 cm long, branched up
to the 4th order, with up to 8(— 12) branches of the
first order, the lower of these in the axils of leaves
or bracts, up to 6 cm long, all rachises sparsely
and shortly appressed -hairy, pedicels up to 5 mm
long, with long appressed hairs. Hypanthium ob-
conoid, 1-2 mm high, sparsely hairy to glabrous
outside. Sepals triangular, 1-1.5 by 1.5-2 mm.
Petals elliptic, 4-4.5 by 2.5-3 mm, white. Sta-
mens c. 20, filaments up to 3 mm, glabrous, an-
thers c. 0.7 mm long. Ovary hairy on exposed top,
2-celled, styles up to 3 mm long, halfway connate,
hairy at base. Fruits obovoid, c. 6 by 4 mm, free
part of hypanthium and sepals closely appressed
against its top, exocarp glabrous, colour unknown,
mesocarp fleshy, endocarp hard and bony. Seeds 3
or 4 per fruit, ellipsoid, 4 by 2 mm, testa with a
hard inner layer and a mucilaginous outer layer,
endosperm thin, cotyledons flat.
Distribution - China, Vietnam; Malesia: Suma-
tra (only seen from Mt Sago), Borneo (only seen
from Mt Kinabalu).
Habitat - In Malesia in primary and secondary
forest, altitude 1 100-1700 m.
5. Photinia serratifolia (Desf.) Kalkman, Blu-
mea 21 (1973) 424. — Crataegus serratifolia
Desf., Catal. Hort. Paris, ed. 3 (1829) 288, 408.
— Type: probably non-existent, see discussion
in Kalkman, I.e.
Photinia serrulata Lindley, Trans. Linn. Soc. Lond.
13 (1821) 103, nom. illeg., superfl.; DC., Prod.
2 (1825) 631; Blume, Bijdr. (1826) 1103; Miq.,
Fl. Ind. Bat. I, 1 (1855) 388; Merr., Enum.
Philipp. Row. PI. 2 (1923) 226; Steenis, Bull.
Jard. Bot. Buitenzorg III, 13 (1934) 242. —
Crataegus glabra auct. non Thunb., Fl. Japon.
(1784) 205: i.a. Aiton, Hort. Kew., ed. 2, 3
(1811) 202; Lindley, I.e., in syn.
Small trees up to c. 15 m. Twigs glabrous, bud
scales up to 13 by 11 mm, dark and hard. Leaves
oblong to elliptic, 8.5-13.5 by 3.5-5.5 cm, base
acute to rounded, margin finely crenate to serrate,
entire only at extreme base, apex acute, sometimes
acuminate, with up to 14 pairs of secondary nerves,
often not distinguishable from stronger tertiary
nerves, venation widely reticulate, not prominent.
316
Flora Malesiana ser. I, Vol. 11 (2) (1993)
with few hairs on both surfaces when very young,
quite glabrous when mature. Petiole 2-3 cm long.
Stipules awn-shaped, 4.5-5.5 by 1 mm, with ex-
centric midrib, early caducous. Inflorescence corym-
bose to semi-globular in shape, up to c. 8 cm long,
with up to 12 first order laterals, these up to 9 cm
long and branched again, the lowermost ones in
axils of leaves, rachises including the 2.5-4 mm
long pedicels glabrous or faintly hairy. Flowers 5-
merous, rarely 4-merous. Hypanthium obconoid,
1.5-2 mm high, upper half free from the ovary,
glabrous outside. Sepals broadly triangular, 1-1.2
by 1.2-1.8 mm. Petals suborbicular to broadly
ovate, 3-4.5 by 3-3.5 mm, white. Stamens 16-
20, filaments up to 3 mm, glabrous, anthers 0.5-
0.8 mm long. Ovary hairy on the free top, 2-, rare-
ly 3-celled, styles up to 2.5 mm long, free. Fruits
subglobular to obovoid with flattened apex, up to
c. 5.5 by 6 mm when dry, upper part of hypan-
thium and sepals closely appressed against top of
fruit, exocarp red (to purple?), mesocarp fleshy,
endocarp rather hard. Seeds 2-4(-6), ellipsoid,
c. 3 mm long, with firm, brown testa, endosperm
a thin layer, cotyledons rather flat.
Distribution - S India, China, Japan, Taiwan;
Malesia: Philippines (seen from Luzon and Minda-
nao) and doubtfully Sumatra. Often cultivated in
Europe.
Habitat - In the Philippines in mossy forest at
c. 2500 m altitude.
Notes - About the name change, necessitated by
Lindley's name being illegitimate, see Kalkman,
I.e.
The one specimen from Sumatra (Jacobs 4363)
is too poor for reliable identification. The descrip-
tion given only pertains to Philippine collections.
PYRACANTHA
Pyracantha Roemer, Fam. Nat. Syn. Monogr. 3 (1847) 104, 219. — Type species: Pyra-
cantha coccinea Roemer (Mespilus pyracantha L,), according to some authors, but not
designated according to Index Nominum Genericorum.
Evergreen shrubs. Twigs often thorny. Carpels 5, free from each other but connate
with the hypanthium, ovaries semi-inferior, styles free, ovules 2. Fruits crowned by per-
sistent sepals, containing 5 pyrenes.
Distribution — Some 12 species in Eurasia. In Malesia one planted species.
Pyracantha angustifolia (Franch.) Schneider,
111. Handb. Laubholzk. 1 (1906) 761. — Coto-
neaster angustifolius Franchet, PI. Delav. (1889)
221. — Types: Delavay 47, 61, 3730, China.
Shrubs, c. 3 m high. Twigs glabrous, some
transformed into thorns. Leaves on short shoots,
oblong, 1-3.5 by 0.5-1 cm, base acute to rounded,
margin enure, apex rounded, often retuse, mucro-
nate, nerves and veins reticulate, hardly visible,
coriaceous, when mature glabrous except remnants
of indumentum near midrib above. Flowers in
short racemes at apex of short shoots, pedicels
hairy, up to 8 mm long. Hypanthium low-campa-
nulate, c. 1 mm high, hairy outside, above the
ovaries with a free rim lined inside by a disc. Sepals
triangular, 1 by 2 mm. Petals orbicular, 3 by 3
mm, spreading in anthesis, white. Stamens c. 20,
filaments up to 2 mm, glabrous, anthers 0.5 mm
long. Ovaries connate with hypanthium over half
their length, hairy on top and dorsally, styles 2 mm,
glabrous. Fruits depressed globular, 3 by 5 mm,
exocarp glabrous, orange (to red?), mesocarp thin-
fleshy, 5 pyrenes remaining free from each other,
their upper parts exposed, endocarps woody. Seeds
2 per cell, flat.
Distribution - Originating from Yunnan, China,
planted as an ornamental in Europe and N America.
Three herbarium specimens seen, collected (in
1953 and 1958) from planted bush(es) along the
Baguio-Bontoc Road, Benguet, Mountain Prov.,
Luzon.
Habitat - Roadside, at an altitude of 1000 (or
2000?) m.
Kalkman — Rosaceae 317
PYRUS
Pyriis L., Sp. PI. (1753) 479. — Type species: Pyrus communis L.
Trees or shrubs, unarmed or with thorns, deciduous. Leaves simple. Flowers in few-
flowered, simple racemes. Hypanthium with a free rim above the ovary, the rim persistent
or falling with the sepals. Ovary inferior, carpels completely adnate with the hypanthium
at base connate with each other, 5-celled, styles free. Fruit a pome, mesocarp fleshy and
with stone cells, endocarp cartilaginous.
Distribution — In Eurasia c. 12 species.
Note — Pyrus communis L., the European pear, is not or hardly cultivated in Malesia.
The Oriental pear, P. pyrifolia (Burm.) Nakai (syn. P. serotina Rehder) is cultivated in
some places and is a promising product. See L.P.A.Oyen in E.W.M. Verheij & R.E.
Coronel (eds.), Edible fruits and nuts, Plant Res. SE Asia (PROSEA Handbook) 2 (1991)
272-276.
RHAPHIOLEPIS
Rhaphiolepis Lindley ex Ker in Edw., Bot. Reg. 6 (1820) t. 468, nom. et orthogr. cons.;
Nakai, J. Arnold Arbor. 5 (1924) 61. — Type species: Rhaphiolepis indica (L.) Lind-
ley ex Ker.
Unarmed small trees or shrubs, evergreen. Leaves simple, margins entire or incised,
nerves not terminating in the margin. Stipules free. Inflorescence a terminal, compound,
rarely simple raceme. Flowers bisexual, 5-merous. Hypanthium obconoid, elongate above
the ovary, the free part at inside covered by a disk. Sepals and upper part of hypanthium
caducous after anthesis. Petals clawed. Stamens 15-20. Ovary inferior, top covered by
the glabrous disk, 2-celled, styles 2, free or connate at base, ovules 2 per cell. Fruit a
pome, globular to (ob)ovoid, with a distinct circular scar at top, mesocarp fleshy, thin,
endocarp thin. Seeds 1 or 2 per fruit, large, testa thin and firm, endosperm absent, co-
tyledons thick. — Figs. 14 d, 15 d.
Distribution — Few species in SE and E Asia, two of them often cultivated as orna-
mentals. In Malesia one wild species and a cultivated one.
KEY TO THE SPECIES
la. Leaves densely woolly when young, glabrate, with finely reticulate darker venation
which is distincdy visible underneath. Hypanthium up to 2.5 mm long, less than half
of it free from the ovary 2. R. philippinensis
b. Leaves only slightly hairy when young, soon quite glabrous, with a more coarse
venation, not darker and not so clearly visible. Hypanthium 3 mm or longer, half of it
or even more free from the ovary 1. R. indica
318
Flora Malesiana ser. I, Vol. 11 (2) (1993)
1. Rhaphiolepis indica (L.) Lindley ex Ker in
Edw., Bot. Reg. 6 (1820) t. 468. — Crataegus
indica L., Sp. PI. (1753) 477. — Type: LINN
sheet 643.11.
Shrubs or small trees. Leaves more or less
crowded, oblong to obovate-oblong, 3-7 by 1-
2.5 cm, base gradually narrowed, margin serrate,
apex acute to obtuse and acuminate, with c. 5 pairs
of nerves, herbaceous to subcoriaceous, with some
hairs when very young, soon glabrous. Petiole up
to 0.5 cm long. Stipules small, caducous. Ra-
cemes compound, up to 13 cm long, the peduncle
very short, rachises practically glabrous, pedicels
up to 5 mm long. Flowers fragrant. Hypanthium
3-3.5 mm high, sparsely hairy outside. Sepals
pointed-triangular, up to 5 by 1-1.5 mm long.
Petals up to 6 mm long, usually white, sometimes
pinkish. Ovary glabrous, styles loosely connate
at base. Fruits globular, c. 6 mm diam., black
when ripe.
Distribution - SE Asia from Thailand to S
China, also in Taiwan and Hainan. The species is
cultivated as an ornamental in many countries, also
in Malesia.
Habitat - On the continent in open types of ever-
green forest, up to c. 1300 m altitude.
Note - The description given is based on speci-
mens cultivated in Java. The variation in the wild
is much larger. See also the note under the follow-
ing species.
2. Rhaphiolepis philippinensis (Vidal) Kalk-
man, Blumea 21 (1973) 434. — Eriobotrya phi-
lippinensis Vidal, Rev. PI. Vase. Filip. (1886)
123; Merr., Enum. Philipp. Flow. PI. 2 (1923)
226; Vidal, Adansonia 5 (1965) 577, in obs. —
Types: Vidal 1350, 1353, both Luzon.
Photinia luzoniensis Merr., Philipp. Bur. Gov. Lab.
Pub. 17 (1904) 18; Philipp. J. Sc, Suppl. 1
(1906) 60; Enum. Philipp. Flow. PL 2 (1923)
226; Steenis, Bull. Jard. Bot. Buitenzorg III, 13
(1934) 242. — Types: Merrill 3223, holo; 3714,
para; both Luzon.
Eriobotrya acuminatissima Nakai, J. Arnold Arbor.
5 (1924) 71. — Type: Martelino & Edaho BS
35622, Panay.
Shrubs or small trees up to 10(-18) m, bark
flaky. Twigs densely woolly when young, glabres-
cent. Leaves oblong or obovate-oblong to lanceo-
late, rarely elliptic, 5-17 by 2.5-5 cm, base
tapering or more rounded, margin sometimes en-
tire, mostly variously incised, at least in the upper
part, apex acute or rounded and acuminate, coriace-
ous, with 7-12 pairs of nerves, often a smaller
nerve of about the same strength between two pri-
mary nerves, venation finely reticulate, woolly
hairy when young on lower surface and near midrib
above, indumentum (almost) disappearing with
age. Petiole 0.5-2.5 cm long. Stipules triangular,
2.5-4 by 0.7-1.5 mm, usually rather long persis-
tent. Inflorescences 2-7.5 cm long, with up to 8
branches, the lower ones often in the axils of
leaves, up to 6 cm long, all rachises including the
2-10 mm long pedicels densely woolly. Hypan-
thium obconoid, 2-2.5 mm high, upper rim free,
densely hairy outside. Sepals triangular, 1.5-3.5
by 1.2-2 mm. Petals elliptic to suborbicular, up
to 6.5 by 4.5 mm, white or whitish. Stamens 16-
20, filaments up to 3.5 mm, glabrous, anthers
0.5-0.8 mm long. Ovary 2-celled, styles free or
shortly cohering, up to 4 mm long, sometimes
glabrous but usually with some hairs at base.
Fruits ovoid to subglobular, 6-9 by 4-9 mm
when dry, with a more or less flattened top but
sepals and hypanthium rim sometimes dropping
off late, exocarp hairy to glabrous, dark purple
when ripe, the mesocarp thin and fleshy, stony
when dry, the endocarp thin. Seeds 1 or 2 per fruit,
rather large, with thin but firm testa. - Figs. 14 d,
15 d.
Distribution - Malesia: Philippines (several is-
lands, also on Palawan), Borneo (only seen from
Sabah).
Habitat - In forest, found at 300-2600 m alti-
tude.
Ecology - Several specimens seen were col-
lected on ultramafic soil. For some other collec-
tions bearing no notes this may also be the case.
Notes - It is certainly not impossible that a
future monographer will decide that this species
has to be included in Rhaphiolepis indica. The
differences are vegetative and unimpressive but that
holds for all species recognized in the genus up till
now.
The leaves of the Philippine specimens are aver-
agely shorter than those in Sabah, but the overlap
is large: Philippines 5-14 by 2.5-5 cm, Borneo
11-17 by 3.5-5 cm.
Kalkman — Rosaceae 319
TRIBUS pruneae
Woody plants with simple leaves. Stipules on the twigs. Epicalyx absent. Pistil 1, supe-
rior, enclosed in hypanthium. Ovules 2, pendulous. Drupaceous fruits, mesocarp some-
times dry. x = 8.
PRUNUS
Prunus L., Sp. PI. (1753) 473; Kalkman, Blumea 13 (1965) 1-115.
Trees or shrubs, rarely with thorns. Buds protected by budscales or naked. Leaves
simple, pinnately nerved, margin incised or entire, with glands in the margin and/or on
the underside or on the petiole. Stipules free or (rarely) connate, on the twigs. Inflores-
cence basically a raceme, rarely branched, in a large part of the genus reduced to a few-
flowered umbel or to only one or two flowers. Flowers normally 5-merous, usually bi-
sexual. Sepals and petals well distinct except in sect. Mesopygeum where the perianth
segments are (sub)equal or irregularly differentiated without (much) difference in size.
Petals usually white or pink. Stamens many (up to 85). Pistil 1, at the bottom of the cup-,
bell- or funnel-shaped hypanthium, at its base often with hairs implanted on the hypan-
thium, also when the ovary itself is glabrous; ovary superior, 1-locular; style terminal,
stigma capitate; ovules 2, pendulous, only 1 normally developing. Fruit a drupe, meso-
carp in wild species not very thick, fleshy to (rather) dry, endocarp bony to woody, thin
to thick Seed with thin testa, without endosperm. — Figs. 18-20.
Distribution — At least 200 species, cosmopolitan. In Malesia c. 35 species and one or
two rarely cultivated ones.
Habitat — The majority of the species is formed by medium (up to 15 m high) or large
(up to 35 m, rarely higher) trees, in different types of forest: primary lowland forest,
montane forest, mossy forest. A good number of species are also found in montane or
subalpine shrubland and then they often are (large) shrubs.
About 44% of the species only occur from sea-level to c. 1500 m altitude, about 32%
only above 1000 m. The remaining 24% of the species can be found as well below 1000 m
as above 1500 m. Only some four species have regularly been collected from altitudes
surpassing 3000 m, going up to c. 3700 m.
Taxonomy — The genus Prunus contains a fair number of useful species with edible
fruits that have since long been domesticated, cultivated, and changed by man. Taxonomy
has in the past often over-classified such groups, giving generic status to each of the culti-
vated species. This has also happenened in Prunus, where apricot, cherry, almond, peach
have been placed in Armeniaca, Cerasus, Amygdalus, Persica, respectively. It is more in
agreement with standards set in the classification of 'useless' groups to unite these genera
and recognize subgenera for some of them. A useful classification of the genus is:
Subgenus Prunus
(among others P. armeniaca L., apricot; P. domestica L., European plum; P. sali-
cina Lindl., Japanese plum)
320 Rora Malesiana ser. I, Vol. 11 (2) (1993)
Subgenus Amygdalus (L.) Focke
(among others P. amygdalus Batsch, almond; P. persica (L.) Batsch, peach and
nectarine)
Subgenus Cerasus (Miller) Focke
(among others P. avium L., sweet cherry; P. cerasus, sour cherry)
Subgenus Padus (Miller) Focke
(among others P. padus; P. serotina)
Subgenus Laurocerasus (Tourn. ex Duhamel) RehcL
The order in which the subgenera are placed, is not phylogenetical, Padus probably
being the most 'primitive' subgenus.
Of the subgenera, only Laurocerasus is represented in Malesia by native wild species,
Padus has one insufficiently known species which may or may not be wild (see Prunus Q,
the other three subgenera are distinctly temperate and not successful in cultivation in the
Malesian region. Only P. persica has been mentioned at the end of the present treatment.
KEY TO FLOWERING SPECIMENS
Specimens bearing flowers only are often not identifiable except by comparing the vegeta-
tive parts with fruiting material.
la. Leaves densely dark-dotted on underside. Stipules intrapetiolarly connate 2
b. Leaves not densely dark-dotted on underside, sometimes pitted on the underside where
hairs have been. Stipules connate or free 3
2a. Leaves without basal glands. Petals 6-7.5 mm long 3. P. mirabilis
b. Leaves with 2 basal glands, usually on petiole, rarely in margin. Petals up to 4 mm
long 2. P. javanica
3a. Basal glands on the petiole 4
b. Basal glands on the leaf surface, sometimes in a contraction of the leaf-base, some-
times absent 5
4a. Racemes without leaves on the basal part of the rachis 1. P. adenopoda
b. Racemes with 2 leaves under the flowers Prunus C
5a. Leaves with 2 or more basal glands and with usually many additional glands in two
rows parallel with the midrib. Petals 3-8 times as long as sepals . 4. P. wallichii
b. Additional glands, when present, not distinctly in two rows parallel with the midrib.
Petals at most twice as long as sepals 6
6a. Basal leaf-glands deeply hollowed, distinctly bulging above 7
b. Basal leaf-glands flat or only slightly hollowed and hardly bulging above, or in all
leaves absent 14
7a. Racemes solitary and simple. Basal leaf-glands in blade proper 8
b. Racemes in bundles (short shoots without or with terminal bud) or truly compound.
Basal glands sometimes in contraction of the leaf-base 12
8a. Stipules free 9
b. Stipules with the bases of their midribs intrapetiolarly connate 11
Kalkman — Rosaceae 321
9a. Stamens 50 or more 23. P. polystachya; 26. P. rubiginosa
b. Stamens not more than c. 50 10
10a. Ovary densely hairy 20. P. marsupialis
b. Ovary glabrous or with few hairs 12. P. fragrans
11a. Stamens 20-40 9. P. dementis
b. Stamens 50-85 23. P. polystachya
12a. Stamens 50 or more. Inflorescences 3.5-1 1 cm long
19. P. malayana; 23. P. polystachya
b. Stamens not more than 50 13
13a. Leaves with 5-9 pairs of nerves. Inflorescences up to 1.5(-4) cm long. Ovary
densely hairy 5c. P. arborea var. densa
b. Leaves with 8-12 pairs of nerves. Inflorescences up to 3.5 cm long. Ovary sparse-
ly hairy to glabrous 31. P. turfosa
See also Primus A with inflorescences up to 10 cm long
14a. Racemes in bundles (short shoots with or without terminal bud) or truly com-
pound 15
b. Racemes solitary and simple 19
15a. Stipules intrapetiolarly connate. Inflorescence a compound raceme, 5-10 cm long,
with 1-5 laterals. Stamens 50-80. Ovary glabrous or with few hairs
19. P. malayana
b. Stipules free 16
16a. Ovary glabrous or with few hairs. Basal leaf-glands absent. Stipules usually with one
large, hollowed gland outside. Stamens 15-40 6. P. beccarii
b. Ovary densely hairy 17
17a. Perianth differentiated as triangular sepals and elliptic to obovate petals. Stamens
35-45 33. P. versteeghii
b. Perianth segments subequal 18
18a. Stamens 10-20 22. P. oocarpa
b. Stamens 10-50(-60) 5. P. arborea
19a. Ovary densely hairy 20
b. Ovary glabrous or with few hairs 28
N.B.: Ovary unknown in Prunus D, fruits sparsely hairy.
20a. Stipules intrapetiolarly connate 9. P. dementis
b. Stipules free 21
21a. Racemes short (-1 cm), peduncle almost none, pedicels up to 2 mm 22
b. Racemes and pedicels longer 24
22a. Leaves thin-papyraceous 15c. P. grisea var. tomentosa
b. Leaves coriaceous, stiff 23
23a. Leaves usually with 2 basal glands. Stipules with 1-3 flat or pustular glands on the
outside. Racemes with up to 6 flowers. Flowers small (hypanthium 1.5 mm long,
perianth segments less than 1 mm long) 14. P. glabrifolia
b. Leaves usually without basal glands. Stipules with glands in margin but not on sur-
face. Racemes with up to 10 flowers. Rowers slightly larger (hypanthium 2-3 mm,
perianth segments 1-1.5 mm long) 21. P. oligantha
322 F!ora Malesiana ser. I, Vol. 11 (2) (1993)
24a. Leaves herbaceous or papyraceous 25
b. Leaves coriaceous, often hard and thick
8. P. brassii; 15a. P. grisea var. grisea; 22. P. oocarpa; 24. P. pulgarensis;
25. P. pullei; 27. P. schlechteri; 30. P. subglabra; 32. P. turneriana
25a. Racemes short, never over 5 cm long 26
b. Racemes (at least some) longer than 5 cm
27. P. schlechteri; 29. P. spicata; 32. P. turneriana; 33. P. versteeghii
26a. Leaves thin-herbaceous to papyraceous 15c. P. grisea var. tomentosa
b . Leaves herbaceous 27
27a. Perianth segments subequal, 7-15
17. P. lamponga; P. odorata (Insufficiently known species)
b. Perianth differentiated as sepals and petals 18. P. laxinervis
N.B.: See also the insufficiently known P. odorata.
28a. Leaves herbaceous to papyraceous 29
b. Leaves coriaceous, stiff 34
29a. Racemes up to 3 cm long 30
b. Racemes longer than 3 cm 32
30a. Leaves papyraceous 31
b. Leaves herbaceous
15a. P. grisea var. grisea; 16. P. kinabaluensis; 17. P. lamponga
31a. Perianth segments subequal 15c. P. grisea var. tomentosa
b. Perianth differentiated as sepals and petals, but equal in size
7. P. brachystachya
32a. Leaves hairy when young, indumentum still present when mature. Stipules often
connate 13. P. gazelle-peninsulae
b. Leaves glabrous or practically so when mature 33
33a. Stipules often connate. Racemes up to c. 30 cm long .... 11. P. dolichobotrys
b. Stipules free. Racemes not longer than 10 cm
15a. P. grisea var. grisea; 16. P. kinabaluensis; 34. P. wallaceana
34a. Racemes shorter than 3 cm
15b. P. grisea var. microphylla; 28. P. sclerophylla
b . Racemes longer than 3 cm
10. P. costata; 15a, b. P. grisea var. grisea and var. microphylla;
16. P. kinabaluensis; Prunus B
Flowers only known as remnants in Prunus D from New Guinea.
Not entered in the key the rarely cultivated P. persica.
KEY TO FRUITING SPECIMENS
la. Fruits ovoid or ellipsoid, distinctly longer than wide 2
b. Fruits (sub)globular or transversely ellipsoid to didymous 11
2a. Leaves dark-punctate on underside 3
b. Leaves not dark-punctate on underside 4
Kalkman — Rosaceae 323
3a. Fruits 21-29 by 13-16 mm. Basal leaf-glands absent 3. P. mirabilis
b. Fruits 15-23 by 7-12 mm. Basal leaf-glands 2, on petiole or in blade-margin
2. P. javanica
4a. Basal leaf-glands on the petiole 1. P. adenopoda
b. Basal leaf-glands on the undersurface 5
5a. Leaves with 2 or more basal glands and with usually many additional glands in 2
rows parallel with the midrib 4. P. wallichii
b. Additional leaf- glands, if present, not in two rows parallel with the midrib .... 6
6a. Inflorescence a compound raceme 7
b. Inflorescence a simple, solitary raceme 8
7a. Fruits 8-1 1 by 6-8 mm 22. P. oocarpa
b. Fruits 18-25 by 16-21 mm 19. P. malayana
8a. Fruits not longer than 13 mm, not wider than 1 1 mm 9
b. Fruits at least 14 mm long, at least 10 mm wide 10
N.B.: Fruits 13-21 by 10-15 mm, prominently beaked, in insufficiently known
taxon, see Note to 15c. P. grisea var. tomentosa.
9a. Leaves with acute base, with 5-8 pairs of nerves. Fruits 10-13 by 8-1 1 mm
21. P. oligantha
b. Leaves with rounded to cordate base, with 7-1 1 pairs of nerves. Fruits 8-11 by 6-8
mm 22. P. oocarpa
N.B.: 10-15 pairs of nerves in Prunus B, of which the fruits are insufficiently
known.
10a. Basal leaf-glands deeply hollowed, distinctly bulging above. Fruits 24-30 by 15-
17 mm, not with an apical point or beak 12. P. fragrans
b. Basal leaf-glands flat or only slighdy hollowed. Fruits 13-24 by 10-20 mm, in-
cluded an apical point or beak of 1-4 mm . 17. P. lamponga; see also Prunus D
11a. Basal leaf-glands deeply hollowed, distinctly bulging above 12
b. Basal leaf-glands flat or only slightly hollowed, or in all leaves absent 22
12a. Basal glands in a contraction of the leaf-base 13
b. Basal glands in the leaf-blade proper 15
13a. Fruits 13-21 by 17-27 mm 23. P. polystachya
b. Fruits not longer than 10 mm, not wider than 12 mm 14
N.B.: Fruits insufficiently known in Prunus A.
14a. Leaves with 8-12 pairs of nerves. Fruits subglobular, 7-9 by 8-9 mm
31. P. turfosa
b. Leaves with 5-9 pairs of nerves. Fruits transversely ellipsoid, 6-8.5 by 8-11.5
mm 5c. P. arborea var. densa
15a. Fruits not longer than 10 mm, not wider than 12 mm 16
b. Fruits longer and/or wider 17
16a. Most of the racemes in bundles (short shoots with or without terminal bud)
5c. P. arborea var. densa
b. Racemes always solitary 20. P. marsupialis
17a. Racemes simple and solitary 18
b. Racemes branched or in bundles (short shoots with or without terminal bud). . . 21
324 Flora Malesiana ser. I, Vol. 11 (2) (1993)
18a. Stipules intrapetiolarly connate 9. P. dementis
b. Stipules free 19
19a. Fruits subglobular to obscurely transversely ellipsoid, 16-17 by 16-20 mm
26. P. rubiginosa
b. Fruits transversely ellipsoid, distinctly wider than long 20
20a. Leaves with 5-8 pairs of nerves. Fruits 6-13 by 7.5-15 mm 20. P. marsupialis
b. Leaves with 9-12(-14) pairs of nerves. Fruits 13-21 by 17-27 mm
23. P. polystachya
21a. Fruits ellipsoid to subglobular, 18-25 by 16-21 mm 19. P. malayana
b. Fruits transversely ellipsoid, 13-21 by 17-27 mm 23. P. polystachya
22a. Racemes in bundles (short shoots with or without terminal bud) or branched, some-
times mixed with solitary, simple ones 23
b. Racemes all solitary and simple 26
23a. Fruits less than 12 mm long, less than 17 mm wide 24
b. Fruits more than 15 mm long, more than 16 mm wide 25
24a. Leaves with 2 basal glands. Fruits 5-11.5 by 7-17 mm 5. P. arborea
b. Leaves without basal glands. Fruits 5-7 by 6.5-10 mm 6. P. beccarii
N.B.: Basal glands usually absent in 5a. P. arborea var. robusta, in which fruits are
9-11.5 by 13.5-^17 mm.
25a. Racemes compound (branched). Fruits subglobular (or ellipsoid), 18-25 by 16-21
mm 19. P. malayana
b. Racemes in bundles (and sometimes partly solitary). Fruits transversely ellipsoid to
didymous, 15-19 by 22-28(-30) mm 33. P. versteeghii
26a. Seeds hairy, sometimes only sparsely so or only near hilum or apex 27
b. Seeds entirely glabrous 32
27a. Ovary and fruit glabrous 28
b. Ovary densely hairy, fruit still with hairs 30
28a. Leaves stiff-coriaceous. Fruits transversely ellipsoid, 6-10 by 8-1 1.5 mm
10. P. costata
b. Leaves herbaceous 29
29a. Leaves glabrous. Stipules free. Fruits subglobular, 12-14 by 13-16 mm
16. P. kinabaluensis
b. Leaves more or less densely hairy when young, hairs not quite disappearing when
mature. Stipules often intrapetiolarly connate. Fruits transversely ellipsoid, 8-12 by
11-17 mm 13. P. gazelle-peninsulae
30a. Fruits 17-33 by 18-34 mm, with thick and woody endocarp 32. P. turneriana
b. Fruits (distinctiy) smaller, endocarp not thick and woody 31
31a. Leaves very hard and stiff, 4-8.5 by 2-3.5 cm, with 6-9 pairs of nerves. Fruits
transversely ellipsoid, 6-7.5 by 7-9 mm 8. P. brassii
b. Leaves herbaceous to coriaceous, 6-17(-20) by 2-8(-10) cm, with 6-13 pairs of
nerves. Fruits transversely ellipsoid to subglobular, 9-16 by 9-18 mm
27. P. schlechteri
See also 5. P. arborea, with racemes normally in bundles but sometimes mixed with
solitary ones.
Kalkman — Rosaceae 325
32a. Fruits more than 20 mm wide 33
b. Fruits at most 20 mm wide 34
33a. Fruits compressed subglobular, 17-33 by 18-34 mm, with thick and woody endo-
carp 32. P. turneriana
b. Fruits transversely ellipsoid to didymous, 15-19 by 22-28(-30) mm
33. P. versteeghii
34a. Fruits (sub)globular, not or not much wider than long 35
b. Fruits transversely ellipsoid to didymous, distincdy wider than long 47
35a. Fruits more than 13 mm long 36
b. Fruits up to 15 mm long 39
36a. Fruits 13-24 by 10-20 mm, including a distinct apical point or beak of 1-4 mm
17. P. lamponga
b. Fruits not distinctly beaked or pointed 37
37a. Fruits with thick, woody endocarp 32. P. turneriana
b. Endocarp not thick 38
38a. Leaves sparsely hairy when young, glabrous when mature ... 30. P. subglabra
b. Leaves densely hairy when young, lower surface remaining hairy when mature
24. P. pulgarensis
39a. Stipules intrapetiolarly connate 9. P. dementis
b. Stipules free 40
40a. Racemes usually not longer than 1 cm 41
b. Racemes normally longer than 1 cm 42
41a. Leaves papyraceous 15c. P. grisea var. tomentosa
b. Leaves coriaceous 21. P. oligantha
42a. Leaves herbaceous 43
b. Leaves stiff -coriaceous 46
43a. Ovary densely hairy and fruit still with hairs 44
b. Ovary and fruit glabrous 45
44a. Leaves (rather) densely hairy when young and hairs remaining on underside when
mature 29. P. spicata
b. Leaves glabrous when mature 18. P. laxinervis
N.B.: See also 15a. P. grisea var. grisea, where specimens from Java and Lesser
Sunda Islands rarely have a densely hairy ovary.
45a. Leaves with usually more than 2, large glands, all or partly above the base in the
blade 16. P. kinabaluensis
b. Leaves usually with 2 glands at the base 15a. P. grisea var. grisea
46a. Leaves (almost) glabrous when mature 15. P. grisea
b. Leaves densely hairy when young and usually still hairy on underside when mature
25. P. pullei
47a. Stipules intrapetiolarly connate 48
b. Stipules free 49
48a. Rachis of inflorescence hairy. Fruits 13-14 by 14-17(-20) mm 9. P. dementis
b. Rachis glabrous or sparsely short-hairy. Fruits 8-11.5 by 11-15 mm
11. P. dolichobotrvs
326 Flora Malesiana ser. I, Vol. 11 (2) (1993)
49a. Leaves stiff-coriaceous 50
b. Leaves herbaceous to papyraceous 53
50a. Leaves (almost) glabrous when mature 51
b. Leaves (densely) hairy when young, still hairy on underside when mature .... 52
51a. Stipules with 1-3 flat or pustular glands on the outside 14. P. glabrifolia
b. Stipules not with glands on the outside 15. P. grisea
52a. Remnant of hypanthium under the fruit often enlarged, 1.5-8 mm diameter. Leaves
often with revolute margins, apex obtuse, sometimes retuse 25. P. pullei
b. Hypanthium remnant under the fruit small. Leaves not with revolute margins, apex
acute or shortly acuminate 28. P. sclerophylla
53a. Leaves papyraceous. Inflorescences not exceeding 3 cm in length 54
b. Leaves herbaceous. Inflorescences usually longer 55
54a. Fruits 8-12.5(-13.5) mm wide. West Malesia . . 15c. P. grisea var. tomentosa
b. Fruits 10-19 mm wide. New Guinea, Australia 7. P. brachystachya
55a. Ovary densely hairy and on fruit still hairs left 56
b. Ovary and fruit glabrous 57
56a. Leaves (rather) densely hairy when young, hairs on underside remaining when ma-
ture. Flowers and fruits sessile 29. P. spicata
b. Leaves (practically) glabrous when mature. Pedicels longer than 1 mm
15a. P. grisea var. grisea
57a. Flowers and fruits sessile 7. P. brachystachya
b. Pedicels at least 1 mm long 58
58a. Leaf apex rounded to broadly acuminate 11. P. dolichobotrys
b. Leaf apex gradually tapering to acuminate 59
59a. Leaves with 5-9 pairs of nerves 15a. P. grisea var. grisea
b. Leaves with 8-14 pairs of nerves 34. P. wallaceana
Fruits not seen or only in a too young stage in the following species, mentioned un-
der Insufficiently known: Prunus odorata from Malaya; Prunus A from Borneo;
Prunus B from Sumatra; Prunus C from Luzon.
Not entered in the key the rarely cultivated P. persica.
Subgenus Laurocerasus
Laurocerasus [Tourn., Inst. (1700) 627, t. 245, 'Lauro-cerasus'] ex Duhamel, Traite
Arbres 1 (1755) 345, t. 133. — Prunus subg. Laurocerasus (Tourn. ex Duhamel)
Rehder, Manual Cult. Trees (1927) 478. — Type species: Prunus laurocerasus L.
Pygeum Gaertner, Fruct. Sem. PI. 1 (1788) 218, t. 46; Koehne, Bot. Jahrb. 51 (1913)
177-224; ibid. 52 (1915) 334-345. — Type species: Pygeum zeylanicum Gaertn. =
Prunus ceylanica (Wight) Miq.
Evergreen, rarely deciduous trees and shrubs. Leaves with entire or incised margin,
basal glands either flat and on the underside of the blade, or flat to cushion-shaped and in
the margin, or on the petiole. Inflorescence a raceme, usually simple but sometimes branch-
ed, the racemes sometimes placed in bundles, in axils of extant or fallen leaves or cata-
Kalkman
Rosaceae
327
phylls. Hypanthium circumscissile after anthesis and its basal part persistent under the
fruit. Perianth regular and biseriate or with (sub)equal segments.
Taxonomy — The inclusion of Pygeum in Prunus subg. Laurocerasus was elaborated
and explained in Blumea 13 (1965).
Laurocerasus and Padus were united into one subgenus of Prunus by Koehne, Bot.
Jahrb. 52 (1915) 279-333, but nowadays it is more usual to keep them as separate sub-
genera (if not as genera, see under the genus description).
The subgenus Laurocerasus can be divided into three sections, as explained in 1965:
— section Laurocerasus, also in Malesia;
— section Mesopygeum containing most but not all of the former genus Pygeum, pre-
dominantly Malesian;
— a section of South-, Central-, and some North-American species which does not
yet have a formal name.
Uses — None of the species is a well-recognized source of useful timber, although of
course the wood may be used locally. Medicinal uses of the bark are less often reported
than could be expected from the presence of cyanogenetic glycosides. Culinary uses of the
leaves are rare. In New Guinea bark of several species is used for the manufacturing of
waist-belts, and it seems to be suitable for basket-work too.
Note — The variation in fruit size is in some cases very large, as appears from the de-
scriptions. This recorded variation is partly caused by the fact that measurements were
taken from dried specimens, where it is often impossible to distinguish between full-grown
fruits and the not yet fully developed ones. Especially in fruits with a fleshy mesocarp this
may give significant differences in size. However, this will not be the only explanation
and at least in some species (e.g. P. turneriana) natural variation seems to be large also
within one individual.
Section Laurocerasus
Leaves entire or with incised margin, basal glands on undersurface, in margin, or on peti-
ole. Perianth regularly biseriate, petals by shape and texture distinct from sepals and (1.2-)
2-8 times as long as the latter. Fruits usually longer than wide, sometimes (sub)globular,
rarely transversely ellipsoid.
Distribution — About 14 species in tropical Africa and tropical Asia, and in adjoining
subtropical to cool-temperate regions: Macaronesia, Portugal, SE Europe, N Iran, China,
Japan. In Malesia 4 species, only Prunus javanic a of wide distribution.
1. Prunus adenopoda Koord. & Valeton, Bull.
Inst. Bot. Buitenzorg 2 (1899) 10. — Prunus
macrophylla Sieb. & Zucc. var. adenopoda Vidal,
Adansonia 4 (1964) 145, comb, illeg. — Lauro-
cerasus adenopoda (Koord. & Valeton) Browicz,
Arbor. Korn. 15 (1970) 6. — Types: Koorders
6419, hololecto; Koorders 10014; both Java.
Prunus pseudoadenopoda Koord., Bull. Jard. BoL
Buitenzorg III, 1 (1918) 84, f. 5. — Type: Koor-
ders 40165, Java.
Prunus javanica auct. non (Teijsm. & Binn) Miq.:
Meeuse & Adelb. in Backer, Bekn. Flora Java,
emerg. ed. IV C 2 (1943) fam. 116, 24, p.p.;
Backer & Bakh. f., Fl. Java 1 (1964) 521, p.p.
Trees up to 12 m. Twigs glabrous. Leaves ellip-
tic to oblong, 8-17 by 4-6.5 cm, base acute to ±
rounded, margin entire, apex acuminate, coriaceous,
with 7-10 pairs of nerves, these hardly prominent
below, venation hardly visible, both sides glabrous,
328
Flora Malesiana ser. I, Vol. 11 (2) (1993)
basal glands mostly 2 on the petiole, large and
protruding. Petiole 7— 1 0(— 1 2) mm long. Stipules
narrowly triangular, 5-12 by c. 1 mm, free, (al-
most) glabrous. Racemes solitary, in the axils of
extant or (rarely) fallen leaves, up to 2.5 cm, pe-
duncle ± absent, rachis pubescent, some empty
bracts at base of raceme, pedicels up to 2.5 mm,
longer under fruit. Hypanthium 1.5-2 mm high,
(almost) glabrous outside. Sepals triangular, c. 1
mm long, ciliate at apex. Petals orbicular, 2.5-3
mm long. Stamens 25-35, filaments up to 5 mm
long, anthers up to 0.5 mm long. Ovary glabrous,
style up to 4.5 mm. Fruits ellipsoid, base attenu-
ate, apex acute, 19-22 by 10-13 mm, mesocarp
probably thick when fully ripe and living fruits
possibly up to c. 24 by 18 mm, endocarp glabrous
inside. Seed with glabrous testa.
Distribution - Java, from Ujung Kulon in West
to Malang Prov. in East), also on Nusakambangan
Island.
Habitat - At low altitudes, up to 500 m, in for-
est but data scarce, also coastal.
Ecology - Some specimens collected on lime-
containing soil.
2. Primus javanica (Teijsm. & Binn.) Miq.,
Fl. Ind, Bat. I, 1 (1855) 365; Koord. & Valeton,
Ic. Bogor. 2 (1904) 169, t. 140; Koord., Adas
1 (1913) pi. 95; Merr., Enum. Born. Row. PL
(1921) 289; Backer & Bakh.f., Fl. Java 1 (1964)
521, excl. syn. P. adenopoda; Prance & Whit-
more in Tree Fl. Malaya 2 (1973) 338; Cock-
burn, Trees Sabah 2 (1980) 98, f. 26. — Cerasus
javanica Teijsm. & Binn., Natuurk. Tijdschr.
Ned. Indie 2 (1851) 309. — Laurocerasus java-
nica (Teijsm. & Binn.) Browicz, Arbor. Kom.
15 (1970) 6. — Type: Teijsmann s.n., Java.
Prunus junghuhniana Miq., PL Jungh. (1855) 402;
Miq., Fl. Ind, BaL I, 1 (1855) 366; Merr., Enum.
Philipp. Row. PL 2 (1923) 234. — Type: Jung-
huhn s.n., Java.
Prunus martabanica Kurz, For. Fl. Brit. Burma 1
(1877) 434; Ridley, Fl. Mai. Penins. 1 (1922)
672. — Type: Wallich 4902.
Platea oblonga Korth. ex Valeton, CriL Overz.
Olacin. (1886) 252. — Type: Korthals s.n.,
Borneo. See Sleumer, Blumea 17 (1969) 248.
Prunus scortechinii (King) Koehne, Bot. Jahrb. 52
(1915) 297. — Prunus martabanica Kurz var.
scortechinii King, J. As. Soc. Bengal 66, 2
(1897) 286. — Types: King's Coll. 5638, Scor-
techini 1782, Malaya.
Prunus forbesii Koehne, BoL Jahrb. 52 (1915) 297.
— Type: Forbes 2728, Sumatra.
Prunus papuana Koehne, Bot. Jahrb. 52 (1915) 299.
— Types: Schultze 22, 49, New Guinea, prob-
ably lost.
Trees up to 35 m, rarely buttressed, bark smooth,
peeling, red- to darkbrown, with distinct smell.
Twigs glabrous, small cataphylls at base of shoots.
Leaves ovate to oblong-ovate, rarely lanceolate,
8-20 by 3-7.5 cm, base rounded to acute, margin
entire, apex tapering to long-acuminate, surface den-
sely black-punctate beneath, herbaceous to slightly
coriaceous, with 8-12 pairs of nerves, venation
not very distinct, both sides glabrous, basal glands
small, usually 2 on the petiole just below the blade,
sometimes in the margin of the blade. Petiole
0.5— 1(— 1.5) cm. Stipules narrowly triangular to
lanceolate, up to 5 by 1.5 mm, connate with their
excentric keeled midribs. Racemes solitary, in axils
of usually fallen leaves, 2-5 cm, in fruit longer,
peduncle very short, rachis glabrous or pubescent,
some empty bracts at base of raceme, pedicels up
to 5(-7) mm. Flowers 5(-6)-merous. Hypanthium
c. 2 mm high, glabrous outside or with few hairs.
Sepals triangular, 0.5-1 mm long, hairy at apex.
Petals elliptic to orbicular, 2.5-4 mm long, white.
Stamens 25-50, filaments up to 6 mm, anthers
up to 1 mm long. Ovary glabrous, style up to
4.5 mm. Fruits ovoid to ellipsoid, base rounded,
apex acute, 15-23 by 7-12 mm, red when ripe,
the endocarp glabrous inside. Seed with glabrous
testa.
Distribution - Burma, Thailand, Vietnam, S An-
daman I.; Malesia: Sumatra, Peninsular Malaysia,
Borneo, Java (not seen from East), Palawan, Cele-
bes, Bali (dubious, only one specimen with ob-
scure label), Moluccas, New Guinea (only seen
from Irian Jaya, up to the border with Papua New
Guinea).
Habitat - In primary and secondary forest, alti-
tude 0-1500 m.
Uses - Rarely noted. Bark used for rice-bins
(Sabah, Tikau SAN 26474), as vermicide for buf-
falo (Sabah, Kandilis SAN 10323) and as fish poi-
son (Kalimantan, Nooteboom 4449).
Note - The length/width index of the leaves
varies from 2 to 4, but narrow-leaved specimens
(especially collected in Borneo, but by no means
restricted to the island) are not separated by a dis-
continuity from the others.
3. Prunus mirabilis Kalkman, Blumea 13
(1965) 49. — Laurocerasus mirabilis (Kalkman)
Browicz, Arbor. Kom. 15 (1970) 6. — Type:
Chew, Corner & Stainton 1097, Mt Kinabalu,
Sabah.
Kalkman — Rosaceae
329
Trees up to 12 m. Twigs glabrous, with 2 cata-
phylls at base of shoots. Leaves elliptic to elliptic-
oblong, 10-15 by 4-6 cm, base rounded, margin
entire, apex acuminate, dark-punctate below, coria-
ceous; with 9-12 pairs of nerves, often on lower
surface a rather strong parallel vein between two
nerves, both sides glabrous, basal glands absent.
Petiole 1-1.2 cm long. Stipules oblong-ovate,
c. 4 by 1.5-2 mm, connate by their excentric
keels, almost glabrous. Racemes solitary, in axils
of fallen leaves, in fruit up to 14 cm long, pedun-
cle 0.5 cm, rachis sparsely pubescent, pedicels
4-6 mm, growing after anthesis. Flowers 5-6-
merous. Hypanthium c. 4 mm high, glabrous out-
side. Sepals rounded-triangular, 2-3 by 1.7-2 mm,
ciliate. Petals ± orbicular, 6-7.5 by 5-7.5 mm,
(almost) glabrous. Stamens c. 40, filaments 6-10
mm, anthers 1 mm long. Ovary glabrous. Fruits el-
lipsoid, base rounded to tapering, apex acute, 21-29
by 13-16 mm, glabrous, probably red when ripe,
endocarp glabrous inside. Seed with glabrous testa.
Distribution - Only known from few specimens
collected in different places on Mt Kinabalu, Sabah.
Habitat- Few notes, at altitudes 1950-c. 3200
m, probably in forest.
Note - Resembling the wider-leaved forms of
P. javanica, but by its large flowers conspicuously
different
4. Prunus wallichii Steudel, Nomencl., 2nd
ed., 2 (1841) 404; Merr., Contr. Arnold Arbor.
8 (1934) 72; Merr., Brittonia 4 (1941) 88;
Prance & Whitmore in Tree Fl. Malaya 2 (1973)
337. — Prunus acuminata (Wall.) Dietr., Syn.
3 (1843) 42, comb, illeg. non Michx. (1803):
Koehne, Bot. Jahrb. 52 (1915) 296, incl. varie-
ties. — Cerasus acuminata Wall., PI. As. Rar.
2 (1831) 78. — Laurocerasus wallichii (Steudel)
Browicz, Arbor. Kom. 15 (1970) 5. — Type:
Wallich 719.
Shrubs or trees rarely larger than 12 m, bark
smooth or warty, brown. Twigs glabrous, some
small cataphylls at base of shoots. Leaves elliptic
to oblong or ovatish, 7-15 by 2-6 cm, base
mostly acute, margin entire, sometimes in upper
half serr(ul)ate, apex acuminate, often thin-herba-
ceous, with 6-9 pairs of nerves, often distinctly
looped and joined, venation hardly visible, both
sides glabrous, basal glands 2 or more, on the
blade-surface, additional glands usually many, in
two rows ± parallel with the midrib. Petiole 0.2-
1 cm long. Stipules narrowly triangular, 4-6 by
0.7-1 mm, free, glabrous. Racemes solitary or in
fascicles of 2-4, axillary, 2-10 cm, peduncle
short, rachis ± glabrous, pedicels 2-8 mm, ± gla-
brous. Hypanthium 1.5-2 mm high, glabrous out-
side. Sepals triangular, 0.5-0.8 mm long, gla-
brous outside. Petals elliptic, 2-4 mm long, white.
Stamens 10-20(-30), filaments up to 3 mm, an-
thers 0.6-0.8 mm long. Ovary densely to sparse-
ly hairy but usually glabrous on the suture, some-
times reduced, style up to 4 mm. Fruits ovoid to
ellipsoid, base rounded, apex ± acute, 10-18 (-20)
by 6-11 mm when dry, probably up to 22 by 15
mm in living state, glabrous, purplish black, me-
socarp thick and fleshy when ripe, endocarp thin,
glabrous inside. Seed with glabrous testa.
• Distribution - Continental Asia (NE India, Pa-
kistan, China, Burma, Thailand, Laos, Vietnam);
Malesia: Sumatra (West and North Prov.), Penin-
sular Malaysia (only seen from Pahang).
Habitat - Montane and subalpine forest and
thickets, at altitudes (600-)1000-3000(-3600) m.
Uses - Seeds edible (W. Meijer, in litt., 1965).
Section Mesopygeum
Leaves always with entire margin, basal glands, if present, on the undersurface. Perianth
with 5-14 segments which are subequal or more or less distinctly differentiated as sepals
and petals, but the latter at most 1.5(— 2) times as long as the former. Fruits usually trans-
versely ellipsoid or didymous, sometimes (sub)globular, in few species ellipsoid.
Distribution — 33 species in tropical Asia, from India to Solomon Islands, 2 species
also in Australia. In Malesia 30 species.
5. Prunus arborea (Blume) Kalkman, Blumea
13 (1965) 90; Prance & Whitmore in Tree Fl.
Malaya 2 (1973) 338; Cockbum, Trees Sabah 2
(1980) 96, f. 26; Corner, Wayside Trees Malaya,
ed. 3, 2 (1988) 619. — Polydontia arborea Blume,
Bijdr. (1826) 1 105. — Pygeum arboreum (Blume)
Blume, M61. BoL nr. 2 (1855) 11; C. Muell. in
Walp., Ann. 4 (1857) 642; Backer & Bakh.f.,
330
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 18. Prunus arborea (Blume) Kalkman. a. Leafy twig, x 0.7; b. bundles of flowering racemes, x 0.7;
c, d. flower, from outside and halved lengthwise, x 4; e. fruiting racemes, x 0.7 (a: De Monchy 1; b-d:
Koorders 6389; e: Goodenough & Ridley 1561).
Kalkman
Rosaceae
331
Fl. Java 1 (1964) 520, p.p., syn. Pygeum lati-
folium excluded. — Pygeum blumei Teijsm. &
Binn., Cat. Hort. Bog. (1866) 252, nom. superfl.
— Type: Blume 654, Java. Blume, Mel. BoL
was effectively published in 1855, see Van
Steenis, Taxon 35 (1986) 272-285. Pygeum
arboreum, therefore, does not have (Blume) EndL
ex C. Muell. in Walp. as author, but (Blume)
Blume. Endlicher, Gen. PI. (1840) already re-
duced Polydontia to Pygeum, but did not make
the specific combination.
Pygeum parviflorum Teijsm. & Binn., Nat. Tijd.
Ned. Indie 2 (1851) 309; Miq., Fl. Ind. Bat. I, 1
(1855) 361; Koord. & Valeton, Bijdr. Booms.
Java 5 (1900) 350, incl. vars.; Koord., Atlas 1
(1913) pi. 112, 113; Ridley, Fl. Mai. Penins. 1
(1922) 675. — Type: Teijsmann & Binnendijk
s.n., cult. Java, L sheet 908.196-192.
Digaster sumatranus Miq., Sum. (1861) 129, 330. —
Pygeum sumatranum (Miq.) Miq., Sum. (1861)
619. — Types: Teijsmann HB 3968, lecto; Jung-
huhn s.n., L sheet 908.191-907; both Sumatra.
For more complete synonymy, see Kalkman, I.e.
Trees up to 35 m, sometimes buttressed, bark
usually smooth, brown or grey, strongly smelling,
sometimes (var. densa, stipulacea, alticola) shrubs.
Twigs more or less densely hairy, more or less gla-
brate with age. Leaves elliptic to oblong or ovate
to lanceolate, 3-25 by 1.5-13 cm, base acute to
subcordate, apex acute to acuminate, herbaceous to
coriaceous, with 5-16 pairs of nerves, sparsely to
densely pubescent when young, indumentum rapid-
ly disappearing or persistent, basal glands usually
2, flat or slightly to distinctly hollowed, Petiole
(0.2-)0.5-1.5(-2) cm. Stipules variable in shape
and size, free, sometimes with conspicuous glands
outside, sometimes rather persistent. Racemes in
axils of extant or fallen leaves, usually fascicled
but sometimes solitary, sometimes truly com-
pound, 0.5-6(-9) cm long, peduncle short, rachis
(densely) pubescent, pedicels 0-6 mm long, pubes-
cent. Flowers fragrant. Hypanthium 1-3 mm high,
pubescent outside. Perianth segments 5-11, usu-
ally subequal, sometimes recognizable as sepals
and petals, 0.5-1 mm long. Stamens 10-50(-60),
filaments up to 7 mm, often hairy at base, anthers
0.2-1 mm long. Ovary densely hairy, style up to
5.5 mm, hairy at base. Fruits globular (not in
Malesia) or subglobular to transversely ellipsoid or
didymous, 5-11.5 by 7-17 mm, more or less
hairy, from green turning white, red, and ultimate-
ly purple or black, the endocarp glabrous or hairy
inside. Seed with hairy or glabrous testa. - Fig.
18.
Distribution - Continental Asia, throughout
Malesia.
Uses - Timber useful in house building. Bark
once reported to be suitable for making rice con-
tainers (var. robusta, Flores, Schmutz 2803).
Note - This variable species was previously
(Kalkman, I.e.) divided into five varieties of which
one, var. montana (Hook, f.) Kalkman, does not
occur in Malesia. A sixth variety, var. alticola, can
now be added. The varieties are connected by odd
intermediary specimens.
KEY TO THE VARIETIES
la. Seeds hairy 2
b. Seeds glabrous 5
2a. Twigs and leaves pubescent when young but
soon glabrous 3
b. Twigs and leaves long retaining their dense in-
dumentum 4
3a. Fruits 5-10.5 by 7.5-13.5 mm. Flowers
small: hypanthium 1-2 mm high
b. var. arborea
b. Fruits 9-11.5 by 13.5-17 mm. Rowers larger:
hypanthium 2-3 mm high d. var. robusta
4a Leaves elliptic to oblong, rarely ovatish, 10-
22(-25) by 4-9(-13) cm, with usually 10-
13 pairs of nerves .... e. var. stipulacea
b. Leaves ovate to elliptic, 6— 13(— 16) by 2.5-
8.5(— 12) cm, with 8-10 pairs of nerves
a. var. alticola
5a Nerves 5-9 pairs. Racemes 0.5-1.5 cm long
c. var. densa
b. Nerves 8-13 pairs. Racemes longer than 2
cm 4
a. var. alticola Kalkman, Blumea 37 (1993) 378.
— Type: De Wilde c.s. 15994, Sumatra.
Twigs densely hairy when young, tardily gla-
brescent. Leaves ovate to elliptic, 6 — 13(— 16) by
2.5 — 8.5(— 12) cm, hard and stiff, densely hairy
when young and also mature leaves usually still
distinctly hairy on midrib above, on midrib and
nerves below, and on petiole, nerves 8-10 pairs.
basal glands 0-2, flat or slightly hollowed. Stip-
ules elliptic to (broadly) ovate, 3-7 by 2.5-7 mm,
sometimes with one or more glands outside. Ra-
cemes solitary or in bundles of 2-4, (l-)2-4(-6)
cm long. Fruits transversely ellipsoid, rarely sub-
globular, 6-10.5 by 7-14(-16) mm, usually still
(sparsely) hairy. Seed glabrous or with few hairs
on testa.
Distribution - Sumatra, Borneo (seen from Sabah
and Sarawak only), Celebes (one specimen only).
332
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Habitat - Montane forest, also mossy forest,
montane scrub, altitude c. ( 1 000 -) 1500 -3000 m.
Note - See note under var. stipulacea. For a com-
parison with Primus oocarpa see under that species.
b. var. arborea — Kalkman, Blumea 13 (1965) 93.
The synonyms given above under the species per-
tain to the type variety.
Twigs pubescent when young, soon glabrous.
Leaves oblong to ovate, sometimes more lanceo-
late, 6-21 by 2.5-8.5(-10) cm, pubescent when
young but early glabrescent, nerves 7-12 pairs,
basal glands usually 2, flat. Stipules elliptic to
ovate, 1.5— 6(— 8) by 1-4 mm. Racemes in bun-
dles of usually 2-5, (l-)2-6(-9) cm long. Fruits
transversely ellipsoid to didymous, 5-10.5 by
7.5-13.5 mm. Seed with densely hairy testa, rare-
ly more sparsely hairy.
Distribution - Throughout Malesia.
Habitat - Primary (and secondary) forest, altitude
0-1800 m.
c. var. densa (King) Kalkman, Blumea 13 (1965)
100.
Pygeum parviflorum Teijsm. & Binn. var. densum
King, J. As. Soc. Beng. 66, 2 (1897) 292. —
Type: King's Coll. 10753, holo; King's Coll.
6986; both Malaya.
Twigs densely pubescent, glabrescent. Leaves
ovate or elliptic to elliptic-oblong, 3-15 by 1.5-8
(-9) cm, densely pubescent when young, usually
still hairy beneath when mature, nerves 5-9 pairs,
basal glands usually 2, flat or distinctly hollowed
and sometimes situated in the contracted leaf-base.
Stipules ovate to elliptic, 1.5-6 by 1-4 mm,
sometimes with one or more crater-like glands out-
side. Racemes in bundles of 2-4, mixed with soli-
tary ones, rarely compound, 0.5-1.5(-4) cm long.
Fruits transversely ellipsoid, 6-8.5 by 8-11.5
mm. Seed with glabrous testa.
Distribution - Peninsular Thailand; Malesia:
Sumatra, Malaya incl. Penang and Tioman Is.,
Borneo.
Habitat - Lowland and montane forest, altitude
(50-)500-2400 m.
d. var. robusta (Koord. & Valeton) Kalkman,
Blumea 13 (1965) 96.
Pygeum robustum (Koord. & Valeton) Koehne,
Bot. Jahrb. 51 (1913) 198. — Pygeum parvi-
florum Teijsm. & Binn. var. robustum Koord.
& Valeton, Bijdr. Booms. Java 5 (1900) 353.
— Types: Koorders 23039, lecto; 6466, 21598,
all Java.
Twigs pubescent when young, soon glabrous.
Leaves ovate or ovate-lanceolate, 10-19 by 4-7
(-10) cm, pubescent when young, early glabres-
cent, nerves 6-10 pairs, basal glands usually ab-
sent. Stipules ovate to elliptic, 3.5-6 by 1.5-3
(-6) mm, often with inconspicuous flat glands
outside. Racemes usually in bundles, sometimes
compound (branched from the base), mixed with
solitary ones, 3-6 cm long. Fruits transversely
ellipsoid to didymous, 9-11.5 by 13.5-17 mm.
Seed with hairy testa.
Distribution - East Java, Bali, Flores.
Habitat - Forest in periodically (very) dry re-
gions, 800-1800 m altitude.
e. var. stipulacea (King) Kalkman, Blumea 13
(1965) 98, p.p.
Pygeum stipulaceum King, J. As. Soc. Beng. 66,
2 (1897) 287; Ridley, Fl. Mai. Penins. 1 (1922)
673. — Type: King's Coll. 11020, Malaya.
Twigs densely hairy when young, tardily glabres-
cent Leaves elliptic to oblong, sometimes ovatish,
10-22(-25) by 4-9(-13) cm, densely hairy when
young, glabrescent but when mature usually still
distinctly hairy at least on midrib, nerves and peti-
ole, nerves (7-)10-13(-16) pairs, basal glands 0-2,
flat or slighdy hollowed. Stipules (broadly) ovate,
4— 9(— 11) by 2.5-6 mm, sometimes with 1-3
glands outside. Racemes in bundles of 2-6, not
rarely also solitary ones, 2-5(-7) cm long. Fruits
transversely ellipsoid, 6— 8(— 11) by 8-13 mm, still
hairy. Seed with hairy testa, sometimes sparsely so,
rarely glabrous.
Distribution - Sumatra, Bangka, Malaya, Singa-
pore, Borneo.
Habitat - Primary and secondary forest, also in
kerangas and mossy forest types, altitude 0-1500
(-2000) m.
Note - In my 1965 revision (p. 98) a possible
subdivision of var. stipulacea was alluded to, but
not executed. New collections make it possible to
extract the higher altitude specimens as a new vari-
ety, alticola.
6. Prunus beccarii (Ridley) Kalkman, Blumea
13 (1965) 104. — Pygeum beccarii Ridley, Kew
Bull. (1938) 281. — Types: Haviland b.r.o.b.,
holo; Haviland 755, Beccari 3516; all Borneo.
Trees up to 27 m, or shrubs, bark smooth,
brown(ish). Twigs sparsely hairy when young.
Leaves elliptic to elliptic-ovate, (6.5-)8-15(-18)
by (3-)4.5-8(-9) cm, base rounded or acute,
apex rounded or obtuse or shortly acuminate, coria-
Kalkman
Rosaceae
333
ceous, with 4— 7(— 1 1) pairs of nerves, venation in-
conspicuous to invisible, lower side and midrib on
upper side sparsely pubescent when young, indu-
mentum rapidly disappearing, basal glands absent,
often some flat foliar glands along the margin.
Petiole 1-2 cm long. Stipules ovate to ovate-
elliptic, 5-7 by 2-5 mm, free, usually with one
large, hollowed gland outside, ciliate. Racemes
mostly in axils of fallen leaves, in fascicles of 3 or
4, or compound with up to 3 laterals, up to 5.5 cm
long, peduncle short, rachis pubescent, pedicels up
to 2 mm long. Hypanthium 1.5-2 mm high,
hairy outside. Perianth segments subequal, 6-10,
0.5-1 mm long, often distant, usually densely
hairy. Stamens 15-40, filaments up to 3.5 mm,
anthers 0.3-0.5 mm long. Ovary with few hairs
or glabrous, style up to 4.5 mm long. Fruits
transversely ellipsoid, 5-7 by 6.5-10 mm, exo-
carp sparsely hairy or glabrous, black when ripe,
mesocarp thin, endocarp glabrous or with hairs
inside. Seed with glabrous or hairy testa.
Distribution - Sumatra, Borneo.
Habitat - In forest, altitude 0-900 m.
Note - Only few collections seen from Borneo,
and only two sterile collections from Sumatra. The
relatively large, hollowed glands in the stipules are
characteristic. The seeds are glabrous in five of the
fruiting specimens seen (most of them immature),
hairy in only one specimen. Judged from the
fruits, the ovary may sometimes be more densely
short- hairy than seen in the flowering specimens.
7. Prunus brachystachya Kalkman, Blumea
13 (1965) 63. — Type: Henty NGF 10526,
Papua New Guinea.
Usually small trees up to 15(— 26) m, sometimes
with low buttresses, bark grey to brown. Twigs
soon glabrous. Leaves oblong to ovate, 7-17 by
2-8.5 cm, base rounded to more acute, apex acu-
minate, herbaceous to papyraceous, with 7-11 pairs
of nerves, venation inconspicuous, both sides
glabrous (maybe some hairs when young), basal
glands 2, flat. Petiole up to 1 cm long. Stipules
narrowly triangular to linear, 4-5.5 by 0.5-1.5
mm, free, hairy outside. Racemes solitary, in axils
of extant or fallen leaves, (0.5— )1.5 — 3 cm long,
peduncle very short, rachis (sparsely) hairy, pedi-
cels 0-1 mm long. Flowers often (?) unisexual,
4- or 5-merous. Hypanthium 1.5-2.5 mm high,
hairy outside, once noted as orange by the collec-
tor. Perianth segments differentiated as sepals and
petals, but the two almost equal in size. Sepals
triangular to elliptic, c. 1 mm long, densely hairy
outside. Petals elliptic, c. 1 mm long, greenish or
white, hairy. Stamens 15-25, glabrous, filaments
up to 3.5 mm, anthers 0.3-0.5 mm long. Ovary
and pistillodium glabrous. Fruits transversely
ellipsoid to didymous, 9-13 by 10-19 mm, exo-
carp glabrous, red to black when ripe, mesocarp
juicy, endocarp glabrous inside. Seed with gla-
brous testa.
Distribution - Papua New Guinea, Irian Jaya
(one collection just on the border with PNG), Aus-
tralia (Queensland).
Habitat - Rain forest, also on riverbanks, alti-
tude up to 450 m.
Notes - The length/width index of the leaves
varies from 2 to 4, but the transition from wide to
narrow leaves is continuous.
Of the four flowering specimens seen, three had
male flowers, the fourth had the stamens and the
ovary both well-developed.
8. Prunus brassii Kalkman, Blumea 13 (1965)
82. — Type: Brass 22814, Papua New Guinea
Trees up to 25 m, or shrubs, bark pale brown.
Twigs densely hairy when young, glabrescent.
Leaves elliptic to elliptic-oblong, 4-8.5 by 2-3.5
cm, base acute to obtuse, apex obtuse and often re-
tuse, very hard and stiff, with 6-9 pairs of nerves,
nerves and venation impressed above, venation
inconspicuous underneath, both surfaces (rather)
densely short hairy when young, glabrescent, basal
glands usually 2, flat. Petiole 0.5-1.2 cm long.
Stipules narrowly triangular, 2.5-5 by 1-2 mm,
free, hairy outside. Racemes solitary, in axils of
extant or fallen leaves, 1.5-4.5 cm long, peduncle
0-0.5 cm, rachis densely hairy, pedicels 1-4 mm
long. Hypanthium 2-3 mm high, densely hairy
outside. Perianth segments 10-12, more or less dif-
ferentiated as sepals and petals, but the two whorls
not entirely regular, 1-1.5 mm long, sepals slight-
ly shorter than petals and with broader base, dense-
ly hairy outside. Stamens 20-30, filaments up to
3.5 mm long, glabrous or hairy at base, anthers
0.5-0.8 mm long. Ovary densely hairy, style up
to 2.5 mm long. Fruits transversely ellipsoid, 6-
7.5 by 7-9 mm, exocarp hairy, colour unknown,
endocarp sparsely hairy inside. Seed with hairy
testa.
Distribution - Papua New Guinea, seen from
Central and Milne Bay Provinces, and with doubt
from S Highlands Province.
Habitat - Montane forest types, also mossy
forest, and in shrubland, 1900-2250(-2750?) m
altitude.
Note - Differs from Prunus pullei, to which it
seems related, in its hairy seeds.
334
Flora Malesiana ser. I, Vol. 11 (2) (1993)
9. Primus dementis (Merr.) Kalkman, Blumea
13 (1965) 70. — Pygeum dementis Merr.,
Philipp. J. Sc, Bot. (1908) 227; Merr., Enum.
Philipp. Flow. PI. 2 (1923) 232. — Types:
Clemens 760, lecto; 966; both Mindanao.
Pygeum apoense Elmer, Leafl. Philipp. Bot. 5
(May 1913) 1623. — Pygeum apoanum Elmer
[in herb.] ex Koehne, BoL Jahrb. 51 (Dec. 1913)
205, nom. illeg. — Type: Elmer 11729, Min-
danao.
Trees up to 30 m, bark grey, rough (only one
note). Twigs hairy, glabrescent. Leaves oblong,
10-22 by 3.5-9 cm, base rounded, apex gradu-
ally tapering to acuminate, herbaceous, with 9-15
pairs of nerves, venation usually not conspicuous,
both surfaces hairy when young, (almost) glabrous
when mature, basal glands 2-4, distincdy hollowed
to flat. Petiole 0.5-0.8 cm long. Stipules ovate
to narrowly triangular, 5-7.5 by 2-4.5 mm, in-
trapetiolarly connate by their keeled midribs, with
inconspicuous marginal glands. Racemes solitary,
axillary, 3-7 cm long, peduncle short, rachis hairy,
pedicels 1-4 mm long, up to 6 mm under the fruit.
Hypanthium 2-3 mm high, hairy outside. Peri-
anth segments 7-10, equal or subequal, c. 1 mm
long, hairy. Stamens 20-40, filaments up to 4.5
mm, hairy at base, anthers 0.3-0.4 mm long.
Ovary densely to more sparsely hairy, style up to
4.5 mm long. Fruits compressed subglobular to
transversely ellipsoid, 13-14 by 14-17(-20) mm,
exocarp more or less hairy to glabrous, dark red
(information scarce), endocarp glabrous inside.
Seed with glabrous testa.
Distribution - Mindanao, Celebes (N peninsula
and central part).
Habitat - Primary and secondary forest, altitude
200-1050 m. Some of the fieldnotes on specimens
from Celebes indicate a preference for volcanic sand
and ultrabasic soil.
Uses - Used as medicine for headache and skin
eruptions {Frake PNH 361 73, probably referring to
the bark).
Note - Only two from the c. 10 collections seen
were collected in more or less recent times, i.e. af-
ter World War II. The Philippine species Prunus de-
mentis, fragrans, and rubiginosa are poorly known.
10. Prunus jcostata (Hemsley) Kalkman, Blumea
13 (1965) 78. — Pygeum costatum Hemsley,
Kew Bull. 1898 (1901) 98. — Type: Giulianetti
(via McGregor) s.n., Mt Scratchley.
Pygeum papuanum Hemsley, Kew Bull. 1898 (1901)
99. — Type: Giulianetti (via McGregor) s.n.,
Mt Scratchley.
Pygeum rigidum Koehne, Bot. Jahrb. 52 (1915)
339. — Type: Ledermann 11453, Papua New
Guinea, not seen.
Pygeum retusum Merr. & Perry, J. Arnold Arbor.
21 (1940) 195. — Types: Brass 9035, holo;
Brass & Meijer Drees 10428, 10439; all Lake
Habbema.
Pygeum hagenianum Gilli, Ann. Naturhist. Mus.
Wien 83 (1980) 455; Kalkman, Blumea 28
(1982) 168 (reduction). — Types: Dosedla 129B,
holo; 745; both Mt Hagen.
Shrubs or small trees, up to 25 m, buttresses
not reported, bark grey to brown, finely fissured
and lenticelled. Twigs glabrous or sparsely hairy
when young. Leaves elliptic to oblong, 3— 13(— 16)
by 2-7 cm, base usually rounded, apex obtuse or
bluntiy acuminate, often retuse, stiff coriaceous,
6-10(-14) pairs of nerves, venation impressed
above, inconspicuous below, puberulous only
when young, both sides glabrous when mature,
basal glands (0-)2-4, flat Petiole up to 1 cm
long. Stipules oblong, 2.5-9 by 1.2-3.5 mm,
free. Racemes solitary, in axils of extant or fallen
leaves, (4— )5— 12(— 15) cm long, peduncle short,
rachis sparsely hairy, pedicels 1-4 mm long,
longer under the fruit. Hypanthium 2-3(-4) mm
high, sparsely hairy to almost glabrous outside.
Perianth regularly 4- to 6-merous, sometimes ir-
regular, hairy outside. Sepals triangular to ovate,
1-2 mm long. Petals ovate to elliptic, 1-3 mm
long, white. Stamens 20-35, filaments up to 5.5
mm, anthers 0.5-1 mm long. Ovary glabrous,
style up to 4 mm long. Fruits transversely ellip-
soid, 6-10 by 8-11.5 mm, fruiting calyx (i.e.
lower part of hypanthium) saucer- to cup-shaped,
3-5 mm diam., exocarp glabrous, red to purplish,
endocarp usually hairy inside. Seed with hairy testa.
Distribution - Throughout New Guinea.
Habitat - Forest and also more open subalpine
habitats like thickets, altitude 1500-3700 m.
Uses - Bark used for making waistbands Q\k&
other species of the genus).
Notes - The species is very similar to Prunus
grisea var. grisea and differs only in the indumen-
tum of the seeds, which is considered to be an im-
portant, usually decisive character for species delim-
itation. When in flower, the two species can hardly
or not be distinguished. The indumentum of the
hypanthium may give a clue: less hairy in costata.
11. Prunus dolichobotrys (K. Schum. & Lau-
terb.) Kalkman, Blumea 13 (1965) 75. — Py-
geum dolichobotrys K. Schum. & Lauterb., Fl.
Schutzgeb. Sudsee (1901) 340; Nachtr. (1905)
Kalkman
Rosaceae
335
274. — Type: Rodatz & Klink 168, Bismarck
Mts, fragment of holotype in WRSL, neotypi-
ficalion (Schlechter 14700) in Kalkman, I.e.,
was not necessary.
Combretum flavovirens Lauterb., Nova Guinea 8
(1912) 847. — Type: Gjellerup 577, Humboldt
Bay.
Trees up to 30 m, buttresses small or absent,
bark brown, lenticelled, peeling off in thin flakes.
Twigs glabrous, often lenticelled. Leaves elliptic
to ovate, (8-)12-26 by 5-12(-15) cm, base
rounded to subcordate, apex rounded to broadly
acuminate, herbaceous, with 7-14 pairs of nerves,
venation transverse, not conspicuous, both sides
entirely glabrous, basal glands 2-4(-6), flat or
slightly hollowed. Petiole up to 1.5(-2) cm long.
Stipules linear to linguiform, 5.5-7.5 by 1.5-2.2
mm, keeled inside and often intrapetiolarly con-
nate, glabrous, sometimes ciliolate. Racemes soli-
tary, axillary, (3-)5-21(-33) cm long, peduncle
up to 1.5 cm, some empty bracts at the base,
rachis glabrous or sparsely puberulous, pedicels
1.5-5 mm long. Flowers sometimes functionally
male, cream-coloured. Hypanthium 2-3 mm high,
(almost) glabrous outside. Perianth differentiated,
(4-)5-merous. Sepals triangular, 0.8-1.5 mm
long, glabrous except sometimes apex and margin.
Petals elliptic to obovate, 1-2 mm long, usually
more hairy than sepals. Stamens 10-30, filaments
up to 6 mm, anthers 0.8-1.5 mm long. Ovary gla-
brous, style up to 4.5 mm long, pistillodium in
male flowers small. Fruits transversely ellipsoid to
didymous, 8-11.5 by 1 1-15 mm, exocarp glabrous,
black when ripe, mesocarp thin, juicy, endocarp gla-
brous inside. Seed with glabrous testa. - Fig. 19.
Distribution - Throughout New Guinea, includ-
ing the Papuan Islands to the East, and also in the
Bismarck Archipelago (New Britain, New Ireland).
Habitat - Primary and secondary forest, often
on riverbanks, altitude 0-1000(-1800) m.
Uses - Hardly ever reported by collectors. Dorn-
streich 76 (E Sepik Prov.): wood used for smaller
house poles, bark for making eel traps, leaves to
flavour soup or cooked greens.
12. Prunus fragrans (Elmer) Kalkman, Blumea
13 (1965) 74. — Pygeum fragrans Elmer, Leafl.
Philipp. Bot. 2 (1908) 475; 5 (1913) 1622;
Merr., Enum. Philipp. Flow. PI. 2 (1923) 232.
— Type: Elmer 7504, lecto, Luzon.
Parinarium coccineum Elmer, Leafl. Philipp. BoL
2 (1909) 578. — Pygeum coccineum (Elmer)
Elmer, Leafl. Philipp. Bot. 5 (1913) 1621. —
Type: Elmer 9787, Negros.
Pygeum megaphyllum Elmer ex Merr., Philipp. J.
Sc, Bot. 10 (1915) 312. — Type: Ramos BS
14923, Luzon.
Small trees, up to 15 m. Twigs sparsely hairy
when young. Leaves elliptic or elliptic-ovate, more
rarely oblong, (10-)12-19(-23) by 4.5-10(-13)
cm, base usually rounded, sometimes shortly de-
current, rarely acute, apex usually acuminate, her-
baceous to ± coriaceous, with 6-10 pairs of nerves,
sparsely pubescent on both sides, basal glands 2,
hollowed and bulging on upper side, other foliar
glands often also hollowed. Petiole 1-2.5 cm
long. Stipules oblong, 4-8 by 1.5-3 mm, free,
with glandular margin. Racemes solitary, in axils
of extant or fallen leaves, up to 8 cm long, in fruit
up to 14 cm, peduncle short, rachis hairy, pedicels
1.5-4(-5) mm long. Flowers bisexual or (rarely)
male. Hypanthium 2.5-3.5 mm high, densely
hairy outside. Perianth segments 7-13, subequal,
up to 1.5(-2.5) mm long, hairy. Stamens 25-55,
glabrous, filaments up to 7 mm, anthers 0.5-0.7
mm long. Ovary glabrous or with few hairs, style
up to 6.5 mm long, also with some hairs or gla-
brous. Fruits ellipsoid, 24-30 by 15-17 mm,
exocarp glabrous, black when ripe, mesocarp thick
and fleshy, hard when dry, endocarp sparsely hairy
inside. Seed with glabrous testa.
Distribution - Philippines, seen from Camiguin,
Luzon, Mindoro, Panay, and Negros. T.C. Huang,
Checklist in Fl. Taiwan 6 (1979) 63, mentions the
species (as Pygeum megaphyllum) for Taiwan,
which is quite plausible. I saw a sterile collection
(C.E. Chang 3080) from Lanyu (Botel Tobago), a
small island off the SE coast of Taiwan. This spec-
imen may belong to P. fragrans, although identifi-
cation remains a bit doubtful.
Habitat - Primary and secondary forest (few data),
altitude 150-2500 m.
13. Prunus gazelle-peninsulae (Kaneh. &
Hatus.) Kalkman, Blumea 13 (1965) 77. —
Pygeum gazelle-peninsulae Kaneh. & Hatus.,
Bot. Mag. Tokyo 52 (1938) 355, f. 1 {'gazelle-
peninsulum). — Type: Kanehira 3966, New
Britain.
Pygeum platyphyllum K. Schum. in K. Schum.
& Lauterb., Fl. Schutzgeb. Sudsee, Nachtr.
(1905) 273. — Type: Nyman 593, 'Sattelberg',
Papua New Guinea.
Trees up to 37 m, buttresses absent or small
spurs, bark brown to grey-brown, smooth or shal-
lowly fissured, flaking. Twigs hairy, glabresent,
older twigs usually lenticelled. Leaves elliptic to
336
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Kalkman — Rosaceae
337
ovate, rarely oblong, (10-) 12-25 by 6-14 cm,
base rounded to subcordate, apex rounded or shortly
acuminate, herbaceous, with 9— 14(-17) pairs of
nerves, venation transverse, both sides more or
less densely hairy when young and indumentum
not quite disappearing with age, basal glands
mosUy 4, flat to slightly hollowed. Petiole up to
1.5 (-2) cm. Stipules triangular to ovate, 3-6.5
by 1.2-2.5 mm, keeled inside and often intra-
petiolarly connate in the lower half, hairy outside
and on margins. Racemes solitary, in axils of ex-
tant or fallen leaves, 5-16 cm long, peduncle very
short, pedicels usually not longer than 1 mm,
hairy as is the rachis. Flowers bisexual or male.
Hypanthium 2.5-4 mm high, densely hairy out-
side. Perianth regular, 5-merous. Sepals triangular
to linguiform, 1-2 mm long, densely hairy. Petals
elliptic to oblong, 1-2 mm long, also hairy,
cream-coloured. Stamens 18-40, filaments up to
8 mm, glabrous, anthers up to 1.5 mm long. Ovary
glabrous except at very base, style up to 6 mm
long, pistillode in male flowers small to minute.
Fruits transversely ellipsoid, 8-12 by 11-17 mm,
fruiting calyx (basal part of hypanthium) saucer-
shaped, 2-3.5 mm diam., exocarp glabrous, black
when ripe, mesccarp fleshy, endocarp mostly hairy
inside. Seed with hairy testa.
Distribution - Moluccas (one flowering speci-
men seen from Ceram), throughout New Guinea,
and Bismarck Archipelago (New Britain).
Habitat - Primary and secondary forest, altitude
0-1600(-2100) m.
Ecology - "Birds feed upon the fruits" (Sayers
21333).
Note - When fruits are absent, distinguishable
from the very similar Prunus dolichobotrys by the
indumentum of the leaves and on the exterior of
the hypanthium.
14. Prunus glabrifolia Kalkman, Blumea 13
(1965) 64; Prance & Whitmore in Tree Fl. Ma-
laya 2 (1973) 338. — Pygeum brevifolium
Hook.f., Fl. Brit. India 2 (1878) 321; Ridley,
Fl. Mai. Pen. 1 (1922) 676. — Types: Griffith
2051, holo; Lobb s.n.; both Mt Ophir, Malaya
Pygeum scortechinii King, J. As. Soc. Beng. 66,
2 (1897) 290; Ridley, Fl. Mai. Penins. 1 (1922)
674. — Type: Scortechini 357, Perak.
Shrubs or trees up to 15 m. Twigs hairy, soon
glabrescent. Leaves elliptic to oblong or ovatish,
3.5-7.5(-ll) by 1.5-4(-5) cm, base acute to
rounded, apex acute to acuminate, coriaceous, with
4-7 pairs of nerves, venation inconspicuous, both
sides entirely glabrous when mature, basal glands
2, flat. Petiole up to 1 cm long. Stipules oblong
to narrowly triangular, 1.5-4 by 0.5-1.5 mm,
free, with 1-3 flat or pustular glands on the sur-
face (only seen in Malayan specimens) and some-
times (also) with marginal glands. Racemes soli-
tary, axillary, up to 1 cm long, with up to 6
flowers, in fruit up to 3.5 cm, peduncle 0, pedicels
0-2 mm long. Hypanthium 1.5 mm high, hairy
outside. Perianth segments 6-9, subequal, less
than 1 mm long, hairy. Stamens 12-20, filaments
up to 3.5 mm, glabrous or with hairs at base, an-
thers up to 0.7 mm long. Ovary densely hairy at
least in lower part, style up to 2.5 mm, hairy or
glabrous. Fruits transversely ellipsoid, 6-9 by
(7-)8.5-12.5 mm, exocarp sparsely hairy, colour
unknown, endocarp glabrous inside. Seed with
glabrous testa.
Distribution - W Sumatra, Malaya (mountains
in Perak, Selangor, Malacca, Johore), Borneo (Sa-
rawak, dubious specimen from Sabah).
Habitat - Both Sumatran specimens and 2 out
of 5 Bornean specimens come from limestone
hills, no notes about the substrate on the 7 speci-
mens from Malaya. The altitude varies from 650
to 1800 m.
Notes - In 1965 the species was only seen from
Malaya and Sumatra; it has now also been collected
in the Gunung Mulu National Park, Sarawak.
A dubious specimen comes from Mt Kinabalu,
Sabah.
The species differs from the rather similar Pru-
nus grisea var. tomentosa in the stiff and hard (not
papyraceous) leaves.
The epithets brevifolia and scortechinii were used
previously in Prunus.
Fig. 19. Prunus dolichobotrys (K. Schum. & Laut.) Kalkman. a. Twig with young racemes, x 0.7; b. Basal
leaf glands, x 0.7; c, d. stipules, x 1, x 2; e. male flowers, x 0.7; f, g. male flower, from outside and
halved lengthwise, x 4 (t = transition petal- stamen); h, i. pistil from bisexual flower, x 4; j. fruit, x 1:
k. seed, x 1 (a, b, e: Hoogland 3966, c, d: Brass 7308; f, g: Womersley NGF 3700; h-k: Darbyshire &
Hoogland8249).
338
Flora Malesiana ser. I, Vol. 11 (2) (1993)
15. Prunus grisea (Blume) Kalkman, Blumea
13 (1965) 56; Prance & Whitmore in Tree Fl.
Malaya 2 (1973) 337. — Pygeum griseum
Blume, Mel. Bot. nr. 2 (1855) 11; C. Muell.
in Walp., Ann. 4 (1857) 642. — Type: Kuhl &
van Hasselt s.n., L sheet 908.197-535, Java.
Blume, Mel. Bot. was effectively published in
1855, see Van Steenis, Taxon 35 (1986) 272-
285. Pygeum griseum, therefore, does not have
Blume ex C. Mueller in Walpers as its author,
but simply Blume.
Germaria latifolia Presl, Epim. Bot. (1851) 221.
— Pygeum preslii Merr., Philipp. J. Sc, Bot.
3 (1908) 227; Elmer, Leafl. Philipp. Bot. 5
(1913) 1621. — Pygeum latifolium (Presl)
Rehder, Bradl. Bibliogr. 2 (1912) 278, comb,
illeg. — Type: Cuming 1815, Bohol, Philip-
pines.
Pygeum latifolium Miq., PI. Jungh. (1855) 401;
Fl. Ind. Bat. I, 1 (1855) 361; Koord. & Valeton,
Bijdr. Booms. Java 5 (1900) 355, incl. varieties;
Koord., Exk. Fl. Java 2 (1912) 336; Adas 1
(1913) pi. 114, 115. — Type: Junghuhn 107,
Java.
Pygeum preslii Merr. var. vulgare Koehne, Bot.
Jahrb. 51 (1913) 203. — Pygeum vulgare
(Koehne) Merr., Enum. Philipp. Flow. PI. 2
(1923) 234. — Type: Borden FB 1806, Luzon.
Pygeum melanocarpum Merr. & Perry, J. Arnold
Arbor. 21 (1940) 191. — Types: Brass 11532,
holo; Brass 11531, Brass & Versteegh 10480;
all Irian Jaya.
Pygeum arboreum (Blume) Blume, p.p.: Backer &
Bakh.f., Fl. Java 1 (1964) 520.
For more complete synonymy, see Kalkman, I.e.
Trees or shrubs. Twigs hairy but rapidly glabres-
cent. Leaves elliptic to oblong, or ovate to lanceo-
late, 2-20 by 1-9 cm, base rounded or acute, apex
various, usually 5-9 pairs of nerves, venation
inconspicuous to invisible, sparsely pubescent to
glabrous on both sides, basal glands usually 2,
flat. Petiole 0.2-1.5(-2) cm long. Stipules 1.5-8
by 0.2-1.8 mm, free. Racemes solitary, in axils
of extant or fallen leaves, 0.5-6.5(-9) cm long,
peduncle 0-1 cm, pedicels 0-7 mm long. Hypan-
thium 1.5-4 mm high, (sparsely) hairy outside.
Perianth segments 6-13, subequal or more or less
distinctly differentiated, 0.5-2 mm long. Stamens
15-50, filaments up to 6 mm, anthers 0.2-0.8
mm long. Ovary usually glabrous, sometimes with
some hairs, rarely distinctly hairy, style up to 7
mm long. Fruits transversely ellipsoid to globular,
6-13 by 7-16 mm, sometimes pointed or beaked
and larger (see note under var. tomentosa), exocarp
(almost) glabrous, via whitish and red turning
purple and finally black, mesocarp thin, endocarp
glabrous or sparsely hairy inside. Seed with gla-
brous testa.
Distribution - Peninsular Burma, Thailand, Viet-
nam; throughout Malesia, var. grisea more in the
eastern part, var. tomentosa more to the west, var.
microphylla endemic to New Guinea.
KEY TO THE VARIETIES
la. Racemes 0.5-l(-2.5) cm, dense, pedicels 0-2
mm. Leaves thin, papyraceous
c. var. tomentosa
b. Racemes 1.5-6.5(-9) cm, loose, pedicels 1-7
mm. Leaves herbaceous or coriaceous ... 2
2a. All or at least most of the full-grown leaves
shorter than 7 cm . . . b. var. microphylla
b. All or at least most of the full-grown leaves
longer than 9 cm a. var. grisea
a. var. grisea — Kalkman, Blumea 13 (1965) 58.
The synonyms given above under the species
pertain to the type variety.
Trees up to 40 m, bark different shades of brown,
smooth or lenticellate. Leaves elliptic to oblong or
ovate to lanceolate, (5-)9-20 by 2.5-9 cm, apex
long-tapering to acuminate, rarely obtuse, herbace-
ous or coriaceous, basal glands (0-)2(-4). Racemes
1.5-6.5(-9) cm, pedicels 1-7 mm. Hypanthium
2-4 mm high. Perianth segments subequal, some-
times (especially in New Guinea) more or less
regularly divided into sepals and petals, up to 2
mm long. Ovary glabrous, rarely with some hairs,
in Java and Lesser Sunda Islands rarely densely
hairy. Fruits transversely ellipsoid to globular, 6-
13 by 8-16 mm, hypanthium remnant under the
fruits up to 2 mm across, but in New Guinea 3-5
mm.
Distribution - Java, Kangean, Philippines, Cele-
bes, Lesser Sunda Islands, Moluccas, New Guinea.
According to the literature also on Taiwan. In Java
and Palawan this variety overlaps with var. tomen-
tosa, in New Guinea with the endemic var. micro-
phylla.
Habitat - Primary and secondary forest, mostly
between sea-level and 2500 m altitude, but especi-
ally in New Guinea often higher (up to 3400 m
altitude), see below.
Notes - The New Guinean plants differ in sev-
eral respects from the plants in Celebes and the
Philippines: their perianth is sometimes larger and
somewhat regularly divided in sepals and petals
(those differing in shape but hardly in size), the
Kalkman
Rosaceae
339
'calyx' (remnant of hypanthium) under the fruit is
large, and the leaves are generally thick and hard.
In New Guinea the collections come from (370-)
1200-3400 m altitude, collections from the Phi-
lippines often bear no data on altitude, but only
two collections were seen from altitudes higher
than 800 m. Also in Celebes all (rather few) col-
lections come from below 1000 m. However, the
morphological differences are very gradual and
characters overlap very much. When two separate
varieties were distinguished, identification would
usually only rest on geography and altitude.
The shapes and dimensions of the fruits show
geographical variation: in New Guinea fruits are
smallest (up to 9 mm long, up to 12 mm wide),
in Java they are also short but often wider (7-10
by 10-15 mm), in the Philippines fruits are larg-
est (8-14 mm long, 9-16 mm wide), with the
fewer specimens from Celebes and the Moluccas
falling within this range. Some specimens from
Palawan have fruits at the extreme end of the varia-
tion and these specimens (Manalo FB 7424. Rids-
dale 182, 191, Soejarto & Fernando 7381) can only
be distinguished from Prunus kinabaluensis by
their foliar glands, see note under that species.
As mentioned under Prunus costata, that species
and the present variety are much alike, among
other characters by their thick and hard leaves and
by the large fruiting calyx (hypanthium remnant).
Flowering specimens cannot always be identified
with certainty.
b. var. microphylla Kalkman, Blumea 13 (1965)
63. — Type: Sleumer & Vink 14250, Irian J ay a.
Usually small trees (rarely over 20 m) or shrubs,
bark rough, brown to grey. Leaves elliptic to ellip-
tic-oblong, (1.5-)2-6(-8.5) by l-3(-5) cm, apex
obtuse or retuse, stiff-coriaceous, basal glands 2
or 4. Racemes 2-5 cm, pedicels 1-4 mm. Hypan-
thium 1 .5 — 3(— 3.5) mm high. Perianth segments
usually subequal, sometimes regularly divided into
sepals and petals, 0.5-1.5 mm long. Ovary gla-
brous. Fruits transversely ellipsoid to subglobular,
6-9 by 7-10.5 mm, hypanthium remnant under
the fruits sometimes large, 1.5-5 mm.
Distribution - Throughout New Guinea.
Habitat - Montane and subalpine forest, altitude
1400-3660 m.
Uses - One collector (Robbins 845) noted the
use of the bark for belts, also known for other spe-
cies.
Note - This is not just a high altitude form of
var. grisea. Both varieties occur at the same alti-
tudes and within var. microphylla the dimensions
of the leaves are not at all correlated with altitude.
The only differences with var. grisea are in the leaf
dimensions and transitional specimens are few.
c. var. tomentosa (Koord. & Valeton) Kalkman,
Blumea 13 (1965) 60. — Pygeum latifolium
Miq. var. tomentosum Koord. & Valeton, Bijdr.
Booms. Java 5 (1900) 358. — Type: Koorders
22255, lecto, Java.
Pygeum maingayi Hook, f., Fl. Brit. India 2 (1878)
319. — Pygeum lanceolatum Hook. f. var.
maingayi (Hook, f.) Ridley, Fl. Mai. Penins. 1
(1922) 674. — Type: Maingay 625, Malaya.
Pygeum hookerianum King, J. As. Soc. Beng. 66,
2 (1897) 293; Ridley, Fl. Mai. Penins. 1 (1922)
676. — Type: Wray 3969, Malaya.
Small trees (rarely over 15 m) or shrubs, bark
brown. Leaves ovate or elliptic to oblong, 4-14
by 1.5-6 cm, apex (long-)acuminate, papyrace-
ous, basal glands (0-)2(-4). Racemes up to
1(— 2.5) cm long, pedicels 0-2 mm. Hypanthium
1.5 — 2(— 3) mm high. Perianth segments 7-10,
subequal, rarely more or less regularly differen-
tiated as sepals and petals, up to 1.3 mm long.
Ovary usually glabrous, sometimes sparsely hairy,
rarely more densely hairy. Fruits transversely ellip-
soid with obtuse apex, to subglobular with an api-
cal point or beak of up to l(-2) mm, 7-12 by
8— 12.5(— 13.5) mm (and see note below).
Distribution - Peninsular Burma, Peninsular and
SE Thailand, Vietnam; in Malesia: N Sumatra,
throughout Malaya incl. Penang, Singapore, Bor-
neo (only Sabah, Sarawak), C and E Java, Philip-
pines (Palawan only), Celebes (only one specimen
seen from Mt Nokilalaki, 1500-1700 m altitude).
Habitat - Primary and secondary forest, altitude
0-1300(-1650) m.
Note - This variety is normally not difficult to
recognize by its thin leaves and short racemes. A
group of (five) collections from Sarawak, mostly
from riverbanks, stands out by having identical
leaves but quite different fruits: those are compres-
sed globular to ovoid, 13-21 by 10-15 by 8-1 1
mm, with a prominent apical beak of up to 4 mm.
These fruits are rather similar to those of Prunus
lamponga, which has, however, different leaves.
The taxonomic status of this Sarawak group re-
mains unsolved as yet.
See also under the insufficiency known Prunus
odorata (p. 349).
16. Prunus kinabaluensis Kalkman, Blumea
13 (1965) 64. — Type: Clemens 29527, Mt
Kinabalu.
340
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Small trees up to 15 m, bark smooth, pale
green to brown. Twigs glabrous. Leaves elliptic to
elliptic-oblong, 6-12 by 2.5-5.5 cm, base acute
to rounded, apex usually gradually tapering, firm-
herbaceous, with 5-7 pairs of nerves, venation
inconspicuous, both sides glabrous, basal glands
0-6, usually more than 2, large, flat, all or part
of them above the base in the blade. Petiole 0.5-
1.5 cm long. Stipules oblong, free, 3.5-5.5 by
0.8-2 mm, ciliolate and sometimes with margi-
nal glands. Racemes solitary, axillary, 2-6 cm
long, peduncle up to 1 cm, rachis sparsely hairy,
pedicels up to 3 mm long. Hypanthium 2-3 mm
high, densely short-hairy to almost glabrous out-
side. Perianth segments 7-12, not regularly differ-
entiated as sepals and petals, triangular to elliptic
or ovate, 1-2 by 0.7-1 mm, cream-coloured to
white, densely hairy to glabrous. Stamens 24-32,
filaments up to 4 mm, glabrous, anthers up to 0.5
mm long. Ovary glabrous, style up to 4 mm long.
Fruits (sub)globular, 12-14 by 13-16 mm, exo-
carp black when ripe, endocarp glabrous or hairy
inside. Seed with glabrous or sparsely hairy testa.
Distribution - Borneo (only seen from Sabah,
Ranau District), Philippines (only seen from
Luzon).
Habitat - Forest, at altitudes between 1300 and
2400 m.
Note - Since the description of this species in
1965, more specimens have been collected in
Sabah, but it is now also known from two collec-
tions from Luzon. The large foliar glands position-
ed above the base remain a conspicuous character
of P. kinabaluensis and by this character it is dis-
tinguishable from the large-fruited specimens of P.
grisea var. grisea that occur on some Philippine
islands, especially on Palawan.
17. Prunus lamponga (Miq.) Kalkman, Blumea
13 (1965) 66; Prance & Whitmore in Tree Fl.
Malaya 2 (1973) 338. — Pygeum lampongum
Miq., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 212.
— Type: Teijsmann HB 4434, Sumatra.
Pygeum goethartianum Koehne, Bot. Jahrb. 51
(1913) 191. — Type: Korthals s.n., Sumatra.
Pygeum gracilipes Koehne, I.e. 191. — Type:
Korthals 126, Sumatra.
Pygeum coriifolium Ridley, J. Str. Br. Roy. As.
Soc. 75 (1917) 30; Fl. Mai. Penins. 1 (1922)
675. — Type: Ridley 14614, lecto; 14616; both
Malaya.
Small trees, rarely more than 15 m high, bark
smooth, brown or greyish. Twigs sparsely hairy
only when very young. Leaves elliptic to oblong,
7— 18(— 21) by 3-6(-9) cm, base acute to rounded,
margin often somewhat revolute, apex (usually
shortly) acuminate, herbaceous, with 4 — 9(— 1 1)
pairs of nerves, often rather wide apart, venation
usually almost invisible, glabrous on both sides
when mature, puberulous only when very young,
rarely more permanently short-hairy on both sides,
basal glands 2(-4), flat. Petiole up to 1 cm long.
Stipuies linear, 3-7 by 0.5-1 mm, free, some-
times with marginal glands. Racemes solitary, in
axils of extant or fallen leaves, up to 2.5 cm long,
peduncle very short, rachis shortly hairy, pedicels
0-2 mm long. Flowers sometimes male. Hypan-
thium 1.7-3 mm high, sparsely hairy outside.
Perianth segments 7— 12(— 15), subequal or at least
not regularly differentiated as sepals and petals, up
to 1 mm long, (sparsely) hairy outside. Stamens
17-32(-40), filaments up to 4 mm, glabrous,
anthers up to 0.5 mm long. Ovary sparsely to
densely hairy, rarely glabrous, style up to 4 mm
long. Fruits ellipsoid to (compressed) subglobular,
13-24 by 10-20 (by 12-14) mm, the length
including an apical point or more distinct beak of
1-4 mm length, exocarp usually sparsely hairy,
rarely glabrous, rarely densely hairy, colour un-
known, endocarp glabrous inside. Seed with gla-
brous testa.
Distribution - Malaya, Sumatra, Bangka, Bor-
neo.
Habitat - Primary and secondary forest of dif-
ferent types, altitudes from sea-level up to c. 850
(-1500) m.
Notes - See the note under P. grisea var. tomen-
tosa for a comparison with a group of specimens
from Sarawak, which have fruits matching the
present species, but different leaves.
Recently collected specimens have not support-
ed the recognition of a variety with more hairy and
also large leaves, as suggested in Kalkman, I.e.
The transitions to small and glabrous leaves are
gradual.
18. Prunus laxinervis Kalkman, Blumea 13
(1965) 69. — Type: Clemens 28477, Borneo.
Trees up to 18 m. Twigs sparsely hairy. Leaves
elliptic to oblong, 10-18 by 5-8 cm, base acute
to rounded, apex acuminate, herbaceous, with 7-9
pairs of nerves, arcuating at some distance from
the margin, venation inconspicuous, both sides
(almost) glabrous, basal glands 2, flat. Petiole up
to 1.3 cm long, hairy to glabrous. Stipules nar-
rowly triangular, 4.5-7 by 1-2 mm, free. Racemes
solitary, axillary, up to 5 cm long, peduncle very
short, rachis hairy, pedicels 1-3 mm long. Hypan-
Kalkman
Rosaceae
341
thium 1.5-3 mm high, hairy outside. Sepals 4,
triangular, up to 1.5 mm long, hairy outside. Petals
4, ovate, up to 1.5 mm long, sometimes thinner
than sepals, hairy, cream-coloured. Stamens 25-
40, filaments up to 3 mm, glabrous, anthers c. 0.5
mm long. Ovary densely hairy, style up to 4 mm
long. Fruits subglobose, 11-13 by 12-14 mm,
exocarp sparsely hairy, colour unknown, endocarp
glabrous inside. Seed with glabrous testa.
Distribution - Only known from Borneo, Sabah
(several collections from Mt Kinabalu and two
slightly deviating ones from other districts).
Habitat - Forest, 1000-1830 m altitude (also
lower?).
19. Prunus malayana Kalkman, Blumea 13
(1965) 102; Prance & Whitmore in Tree Fl.
Malaya 2 (1973) 338. — Type: Nur SFN 32695,
Malaya.
Trees up to 30 m, buttresses none or short, bark
grey to brown, smooth or lenticellate. Twigs hairy
when young. Leaves elliptic, 12-22 by 6-13 cm,
base truncate to subcordate, apex obtuse to shortly
and bluntly acuminate, herbaceous, with 9-15 pairs
of nerves, venation indistinct, upper side glabrous,
lower side shortly hairy and glabrescent, basal
glands usually 2, large, flat or slightly hollowed
and bulging above, sometimes glands absent. Peti-
ole up to 1.5 cm long. Stipules pointed ovate, 3-5
(-11) by 2-4 mm, connate over up to 1 mm by
their excentric, keeled midribs. Racemes com-
pound, in axils of extant or fallen leaves, 5-10 cm
long, with 1-5 laterals up to c. 7 cm, peduncle
c. 1 cm, rachis hairy, pedicels up to 2.5 mm long.
Flowers sometimes male, sex disposition in trees
unknown. Hypanthium 2-3 mm high, shortly
hairy outside, with long hairs at base inside. Peri-
anth differentiated as sepals and petals, but these
only differing in shape, not in size, 5-6-merous.
Sepals triangular, c. 1 mm long. Petals elliptic,
c. 1 mm long, densely hairy. Stamens 50-80,
filaments up to 4.5 mm, glabrous, anthers c. 0.5
mm long. Ovary glabrous or with some few hairs,
style up to c. 3 mm long, pistillodium in male
flowers minute, hidden in the hairs on the bottom
of the hypanthium. Fruits ellipsoid to (rarely) sub-
globular, 18-25 by 16-21 mm, exocarp glabrous,
colour when ripe unknown, endocarp glabrous or
sparsely hairy inside. Seed with glabrous testa or
with few hairs especially near apex.
Distribution - Peninsular Malaysia: Pahang,
Trengganu, Perak.
Habitat - Forest, primary or disturbed, at alti-
tudes from c. 100 to c. 1200 m.
Note - This species is, by its large leaves and
compound racemes similar to Prunus polystachya,
and is obviously closely related. The main differ-
ence is in the fruits (ellipsoid vs. transversely el-
lipsoid) and in flower the two are not always easy
to distinguish. See also under Insufficiently known
species, Prunus A.
20. Prunus marsupialis Kalkman, Blumea 13
(1965) 71. — Pygeum glandulosum Merr.,
Philipp. J. Sc, Bot, 3 (1908) 226; Elmer,
Leafl. Philipp. Bot. 5 (1913) 1621; Merr.,
Enum. Philipp. Flow. PI. 2 (1923) 233. —
Types: Williams 642, lecto; 25 syntypes; all
Philippines.
Pygeum pubescens Merr., Philipp. J. Sc, Bot. 9
(1914) 359. — Type: Wenzel 333, lecto; 5 para-
types; all Philippines.
Small trees up to 13 m. Twigs hairy when
young. Leaves elliptic to oblong, or somewhat
ovatish, (6-)8-13(-16) by 3-6.5 cm, base round-
ed to acute, apex obtuse to acuminate, herbaceous,
with 5-8 pairs of nerves, venation reticulate to
more or less transverse, usually not conspicuous,
both sides pubescent when young, glabrate, basal
glands normally 2, hollowed and distinctly bulging
above, rarely absent. Petiole up to 1 cm long. Stip-
ules linear, free, often persistent, 4— 12(— 17) by
0.5-3 mm, with marginal glands. Racemes soli-
tary, in axils of extant or fallen leaves, up to 7(-
10.5) cm long, peduncle short, rachis hairy, pedi-
cels up to 1.5(— 3.5) mm long. Hypanthium c. 2
mm high, densely hairy outside. Perianth segments
8-12, subequal, up to 1.5 mm long, hairy. Sta-
mens 20-30(-40), filaments up to 5 mm, gla-
brous, anthers 0.3-0.4 mm long. Ovary densely
hairy, style up to 4 mm long, base hairy. Fruits
transversely ellipsoid, 6-13 by 7.5-15 mm, exo-
carp still sparsely hairy, via red ultimately becom-
ing black, endocarp glabrous or with some hairs
inside. Seed with glabrous testa. - Fig. 20.
Distribution - Philippines, many islands from
north to south, but not (yet) seen from Mindanao
and Palawan. T. -C.Huang, Checklist in Fl. Tai-
wan 6 (1979) 63, mentions the species for Taiwan,
from where I have not seen any specimens.
Habitat - Forest and forest edges, from sea-level
to c. 1100 m altitude.
Note - Both epithets published in Pygeum are
not available in Prunus.
21. Prunus oligantha Kalkman, Blumea 13
(1965) 83. — Type: Hoogland & Pullen 5439,
Papua New Guinea.
342
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 20. Prunus marsupialis Kalkman. a. Flowering twig, x 0.7; b. basal leaf glands, x 2; c, d. flower,
from outside and halved lengthwise, x 4; e. fruits, x 0.7 {Loher 2225).
Kalkman — Rosaceae
343
Small trees, rarely over 15 m high, sometimes
shrubs, bark dark brown or grey. Twigs hairy.
Leaves oblong to ovate-oblong, 4-1 1(-14) by
1-4 cm, base acute, rarely rounded, apex acute to
acuminate, coriaceous, with 5-8 pairs of nerves,
venation invisible, densely hairy when young,
hairs persistent on lower side, basal glands 0(-2),
small, flat. Petiole up to 1 cm long. Stipules nar-
rowly triangular, 2.5-5 by 1-1.5 mm, free, with
marginal glands. Racemes solitary, axillary, up to
1 cm long, in fruit no longer than 2 cm, rachis
densely hairy. Flowers up to 10, sessile, yellow-
ish. Hypanthium 2-3 mm high, hairy outside.
Perianth segments subequal, 6-12, 1-1.5 mm
long. Stamens 12-30, filaments up to 2.5 mm,
glabrous or hairy at base, anthers c. 0.5 mm long.
Ovary densely hairy, style up to 3 mm long, some-
times hairy at base. Fruits ellipsoid or subglobu-
lar, 10-13 by 8-11 mm, exocarp still hairy when
ripe, red, turning black when ripe, endocarp gla-
brous inside. Seed with glabrous testa.
Distribution - New Guinea, only one collection
seen from Irian Jaya.
Habitat - Primary or disturbed montane forest,
at 1500-2900 m altitude.
Uses - The bark is used for making waist-belts
(Clunie & Katik LAE 63312, Vink 16519), as re-
ported for other species too.
22. Prunus oocarpa (Stapf) Kalkman, Blumea
13 (1965) 102. — Pygeum oocarpum Stapf,
Trans. Linn. Soc. Bot. 4 (1894) 144; Merr.,
Enum. Born. Flow. PI. (1921) 289. — Type:
Haviland 1118, Mt Kinabalu.
Shrubs or trees up to 15 m. Twigs densely
hairy when young and rather tardily glabrescent.
Leaves ovate to elliptic-ovate, 4— 10(— 13) by 3-6
(-8) cm, base rounded to cordate, sometimes de-
current, margin often revolute, apex rounded to
obtuse, thick and coriaceous, with 7-11 pairs of
nerves, venation slightly impressed above, flat
underneath, densely hairy when very young, hairs
disappearing with age, leaving the surfaces distinct-
ly pitted and midrib above usually still hairy, basal
glands 2(-4), flat to slightly hollowed and bulging
above. Petiole up to 0.5 cm long, rarely longer.
Stipules triangular, 3-5 by 2-3 mm, free. Ra-
cemes solitary, simple or with l(-2) side-branch(es)
at very base, in axils of extant or (more rarely)
fallen leaves, up to 3(-5) cm long, peduncle ab-
sent, rachis hairy, pedicels up to 2 mm long.
Hypanthium 1.5-2 mm high, densely and rather
long hairy outside. Perianth segments (sub)equal,
7-1 1, 1-2 by 0.3-0.7 mm, densely hairy, brown-
ish to yellowish. Stamens 1 1— 18(— 21), filaments
up to 2.5 mm, anthers 0.3-0.5 mm long. Ovary
densely long-hairy, style 2-3 mm long, some-
times hairy at base. Fruits ellipsoid, 8-11 by
6-8 mm, hairy, red when ripe (?), endocarp gla-
brous inside. Seed with glabrous testa.
Distribution - Borneo, known from Sabah (Mt
Kinabalu, Mt Trusmadi), Brunei (Mt Pagon Periok),
and Sarawak (Mt Api).
Habitat - Mossy forest and other types of for-
est, often on mountain ridges, also in shrubland,
reported from limestone as well as from ultramafic
soil and from sandstone, alL 1200-3000 m.
Note - This species is quite homogeneous in
appearance. It resembles some specimens of P. ar-
borea var. alticola, but differs in the shape of the
fruits: transversely ellipsoid and nearly always
broader than long, rarely subglobular, in var. alti-
cola, ellipsoid and always longer than wide in P.
oocarpa. Flowering specimens are sometimes more
difficult to distinguish, but var. alticola has more
and often longer stamens (16-40, filaments up to
6 mm). The crater- like glands that sometimes occur
on the stipules of var. alticola, were not seen in P.
oocarpa.
23. Prunus polystachya (Hook, f.) Kalkman,
Blumea 13 (1965) 88; Prance & Whitmore in
Tree Fl. Malaya 2 (1973) 339; Corner, Wayside
Trees of Malaya ed. 3, 2 (1988) 619, pi. 199.
— Pygeum polystachyum Hook, f., Fl. Brit.
India 2 (1878) 320; Ridley, Fl. Mai. Penins. 1
(1922) 674. — Type: Maingay 627, Malacca.
Pygeum myriandrum Merr., Pap. Mich. Acad. Sc.
19 (1934) 155. — Type: Bartlett 6871, Sumatra.
Deciduous trees, up to 35 m high and more than
60 cm diam., sometimes with buttresses, bark
grey to black or blackish brown. Twigs hairy only
when young. Leaves elliptic to elliptic-ovate, 8-26
by 5-15 cm, base rounded to truncate, rarely acute,
apex obtuse or shortly acuminate, herbaceous to
coriaceous, with 9— 12(— 14) pairs of nerves, vena-
tion inconspicuous, glabrous on both sides, finely
pubescent below only when very young, basal
glands (0-)2, deeply hollowed, distinctly bulging
above, not rarely in the contracted leaf-base. Petiole
0.5-1.5 cm long. Stipules narrowly triangular to
oblong, oblique, 4-10 by 1.5-3 mm, usually
free, rarely shortly intrapetiolarly connate. Racemes
solitary, mainly in the basal pan of shoots and ap-
pearing with the flush of new leaves, in the axils
of leaves or in kataphylls below the leaves, or
united into pseudo-panicles (leafless shoots usually
retaining their terminal bud) or fascicled by contrac-
344
Flora Malesiana ser. I, Vol. 11 (2) (1993)
tion of the main axis, or truly compound with
some side-branches and without terminal bud,
3.5-11 cm long, rachises short-hairy, pedicels 1-3
mm long. Flowers sometimes male, fragrant. Hy-
panthium 2-3 mm high, shortly hairy outside.
Perianth segments 7-11, subequal or inequal but
only rarely by their shape somewhat regularly dif-
ferentiated as sepals and petals, 1-1.5 mm long,
hairy. Stamens 50-85, filaments up to 5 mm,
glabrous, anthers 0.3-0.7 mm long. Ovary gla-
brous except around insertion, or with some long
hairs higher up, often on one side only, rarely
more densely hairy, style 4-5 mm long, pistillo-
dium in male flowers minute. Fruits transversely
ellipsoid, 13-21 by 17-27 mm, exocarp glabrous,
remaining green when mature (?), endocarp gla-
brous inside. Seed with glabrous or hairy testa.
Distribution - Sumatra (only seen from the
middle part), Malaya, Singapore, Borneo? (From
this island only one collection was seen, viz. For-
man & Blewett 946, Brunei; it has the typical
large leaves and fruits of this species but deviates
in having flat instead of marsupial basal leaf glands).
Habitat - Primary and secondary forest, altitude
0-600 m.
Ecology - Corner, I.e., records pollination by
"crowds of hover-flies and small beetles", attracted
by the fragrant flowers. He invented the queer
English name "bat's laurel" for the species but the
role of bats was not elucidated.
Notes - See Kalkman, Blumea 13 (1965) 9,
fig. 4, for a discussion of the morphology of the
inflorescence in this species.
Closely related is Primus malayana, see the note
under that species. See also under Insufficiently
known species, Prunus A.
24. Prunus pulgarensis (Elmer) Kalkman,
Blumea 13 (1965) 67. — Pygeum pulgarense
Elmer, Leafl. Philipp. Bot. 5 (1913) 1627;
Merr., Enum. Philipp. Flow. PI. 2 (1923) 234.
— Type: Elmer 13200, Palawan.
Pygeum monticolum Merr., Philipp. J. Sc, Bot.
10 (1915) 312. — Type: Whitford 1203, Luzon.
Small trees. Twigs densely hairy. Leaves ellip-
tic to oblong or ovatish, 7-14 by 3-5.5 cm, base
acute or rounded, apex acuminate, coriaceous, with
6-9 pairs of nerves, venation reticulate, impressed
above, inconspicuous below, densely hairy when
young, lower surface remaining hairy when mature,
basal glands 2-4 or absent, flat. Petiole 0.5-1.2
cm long. Stipules elliptic, c. 3-4.5 by 1-2 mm,
free. Racemes solitary, in axils of extant or fallen
leaves, up to 5.5 cm long, peduncle absent, rachis
densely hairy, pedicels up to 1 mm long. Hypan-
thium 2-2.5 mm high, densely hairy outside.
Perianth segments subequal, 8-10, up to 2 mm
long, densely hairy. Stamens 25-35, filaments up
to 5 mm, glabrous or hairy at base, anthers c. 0.5
mm long. Ovary densely hairy. Fruits globular,
14-18 mm diam., exocarp hairy, mesocarp thick
and hard when dry, endocarp thin, glabrous inside.
Seed with glabrous testa.
Distribution - Philippines (Luzon, Palawan).
Habitat - Mountain forest, c. 1000-1200 m al-
titude.
Note - Little known species and no new mate-
rial added to what was known in 1965.
25. Prunus pullei (Koehne) Kalkman, Blumea
13 (1965) 85. — Pygeum pullei Koehne, Bot.
Jahrb. 52 (1915) 338. — Type: Pulle 1005,
Irian Jaya.
Prunus pullei (Koehne) Kalkman var. grandiflora
Kalkman, Blumea 13 (1965) 86. — Type: Pul-
len 252, Papua New Guinea.
Small trees, up to 15(-24) m, in higher alti-
tudes usually treelets of some metres or large shrubs,
bark brown, usually rough and lenticellate, some-
times grey. Twigs densely hairy, Leaves elliptic to
oblong, 2-12 by 1.5-5 cm, base acute to round-
ed, margins often revolute also when living, apex
obtuse, often retuse, stiff-coriaceous, with 5-9
pairs of nerves, distinctly looped and joined near
the margin, densely hairy when young and usually
still hairy below when mature, basal glands 2(-4),
flat. Petiole 0.2-1 cm long. Stipules narrowly
triangular, 2.5-7 by 0.7-1.8 mm, free. Racemes
solitary, in axils of extant or fallen leaves, 1-12
cm long, peduncle 0-1.5 cm, rachis densely hairy,
pedicels 0-7 mm long. Hypanthium 2-4 mm
high, densely hairy outside. Perianth segments 8-
12, subequal or unequal but not regularly differen-
tiated as sepals and petals, 1-2 mm long, hairy
outside. Stamens 15-40, filaments up to 7 mm,
glabrous or with some hairs at base, anthers 0.4-1
mm long. Ovary densely hairy, style up to 5 mm
long, sometimes hairy at base. Fruits subglobular
to transversely ellipsoid, 6-11 by 7-11.5 mm,
exocarp hairy, shining purplish black when ripe,
endocarp glabrous or with some hairs inside, calyx
(i.e. remnant of hypanthium) under the fruit 1.5-4
mm diam., but in specimens from high altitudes
up to 8 mm diam. Seed with glabrous testa.
Distribution - New Guinea.
Habitat - All kinds of montane forest, also mos-
sy forest, and in subalpine shrubland, at (1500-)
2000-3700 m altitude.
Kalkman
Rosaceae
345
Uses - As for other species, there are a few re-
ports {Bowers 843. Pullen 252) of the use of the
bark for making men's waist belts.
Notes - Herbarium material has more than doub-
led since 1965 and it has become clear that the ear-
lier distinction of two varieties (Kalkman, I.e.)
cannot be upheld. Leaf and flower characters are
not really correlated and intermediates occur fre-
quently. The infraspecific variation of course re-
mains. Large leaves (more than 8 cm in length)
are only found in lower altitudes (below 3000 m),
but there is not a real correlation of leaf size and
altitude since small leaves (up to 7 cm long) occur
throughout the altitudinal range. The distinctness
of the nervation underneath depends much on the
size of the leaves: larger leaves often have more
prominent nerves. The size of the flowers, i.e.,
the hypanthium, and correlated with it the size
of the fruiting calyx, is variable but a boundary
can only arbitrarily be drawn. There is a tendency
that larger flowers especially occur in higher alti-
tudes.
26. Prunus rubiginosa (Elmer) Kalkman, Blu-
mea 13 (1965) 72. — Pygeum rubiginosum
Elmer, Leafl. Philipp. Bot. 5 (May 1913) 1624;
Merr., Enum. Philipp. Fl. PI. 2 (1923) 234. —
Type: Elmer 11857, holo; 14067; both Minda-
nao.
Pygeum elmerianum Koehne, Bot. Jahrb. 51 (Dec.
1913) 206; Merr., Enum. Philipp. FI. PI. 2
(1923) 232. — Type: Elmer 12210, Sibuyan.
Trees up to c. 17 m, bark dark brown, smooth
(few reports only). Twigs rapidly glabrescent.
Leaves elliptic to oblong, 7-15.5 by 3-7.5 cm,
base acute or rounded and decurrent, apex gradually
narrowing or subacuminate, herbaceous, with 7-10
pairs of nerves, venation not conspicuous, sparsely
hairy to glabrous, basal glands 1-2, distinctly hol-
lowed and bulging above. Petiole 0.5-1.5 cm long.
Stipules 3-6 by 1-1.8 mm, free, hairy, with mar-
ginal glands. Racemes solitary, in axils of extant
or fallen leaves, 2-7 cm long, peduncle very short,
rachis hairy, pedicels 0-1 mm long. Hypanthium
2.5-3 mm high, hairy outside. Perianth segments
10-12, not regularly differentiated as sepals and
petals, up to 2 mm long. Stamens 60-75, fila-
ments c. 3.5 mm, glabrous, anthers 0.3-0.5 mm
long. Ovary more or less hairy, style c. 5 mm long.
Fruits compressed subglobular to obscurely trans-
versely ellipsoid, 16-17 by 16-20 mm, exocarp
almost glabrous, red turning purple, mesocarp
leathery-fleshy, endocarp glabrous inside. Seed with
glabrous testa.
Distribution - Philippines, known from Luzon,
Mindoro, Sibuyan, and Mindanao, but in each is-
land only from 1 or 2 collections.
Habitat - Forest, altitude 250-1200 m.
Note - Young fruits seem to be ellipsoid.
27. Prunus schlechteri (Koehne) Kalkman,
Blumea 13 (1965) 79. — Pygeum schlechteri
Koehne, Bot. Jahrb. 51 (1913) 210. — Type:
Schlechter 17621, Papua New Guinea.
Pygeum forbesii Koehne, I.e. 210. — Type: Forbes
529, Papua New Guinea.
Pygeum laurocerasus Koehne, I.e. 208. — Type:
Schlechter 18621, Papua New Guinea.
Pygeum tetradenium Koehne, BoL Jahrb. 52 (1915)
341. — Type: Ledermann 7889, Papua New
Guinea.
Pygeum salomonense Merr. & Perry, J. Arnold
Arbor. 21 (1940) 1%. — Type: Brass.2727, San
Crist6bal.
Trees up to 35 m, sometimes buttressed, bark
(dark to greyish) brown, smooth or shallowly fis-
sured and with vertical lines of lenticels. Twigs
sparsely to densely hairy. Leaves elliptic to ob-
long, rarely ovate, 6-17(-20) by 2-8(-10) cm,
base rounded to acute, apex tapering or acuminate,
herbaceous to coriaceous, with 6-13 pairs of
nerves, venation usually inconspicuous, sparsely
to densely hairy when young, upper surface be-
coming (almost) glabrous, lower surface usually
still hairy when mature, basal glands (0-)2-6,
flat. Petiole 0.2-1 cm long. Stipules ovatish to
triangular, 2.5-6(-14) by 1-2.5 mm, free, usu-
ally with marginal glands. Racemes solitary,
mostly in axils of fallen leaves, 1 .5— 7(— 15) cm
long, peduncle short, rachis densely hairy, pedicels
0-2(-4) mm long. Flowers fragrant, sometimes
male. Hypanthium 1.5-3 mm high, hairy out-
side. Perianth segments subequal and not regularly
differentiated as sepals and petals, 7-13, up to 2
mm long. Stamens 15-40, filaments up to 5 mm,
glabrous or hairy at base, anthers 0.5-1 mm long.
Ovary densely hairy, style up to 3 mm long, often
hairy, pistillode in male flowers minute, hairy.
Fruits transversely ellipsoid to subglobular (see
note), in New Guinea and New Ireland 9-16 by
9-18 mm, in the Solomon Islands up to 17.5 mm
long and 20(-23) mm wide, usually hairy, purplish
black when ripe, mesocarp often rather thick, endo-
carp glabrous or with few hairs inside. Seed with
hairy testa, sometimes hairs only near the hilum.
Distribution - New Guinea, Bismarck Archi-
pelago (New Ireland), Solomon Islands (from Bou-
gainville to San Crist6bal).
346
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Habitat - Primary and secondary forest, in New
Guinea from sea-level to c. 2600(-2800) m alti-
tude, in the Solomon Islands not above 1200 m.
Uses - Few reports on medicinal use are known
only from the Solomon Islands (Kajewski 2383,
2483): a macerate of the bark is applied to aching
teeth and sore legs.
Notes - In New Guinea one could distinguish
two groups differing in the shape and size of the
fruits:
I Mature fruits transversely ellipsoid, sometimes
distinctly bilobed, 10-13 by 12-18 mm. Seen
from low altitudes, up to 1220 m.
n Mature fruits (sub)globular, 9-16 by 9-16
mm. Seen from higher altitudes, (1280-)1680-
2620 m.
The two groups overlap in the measurements of
their fruits and the distinction in shape is not al-
ways evident: the (sub)globular fruits of group II
may tend to be transversely ellipsoid. Moreover,
the characters of leaves and flowers do not show
any correlation with the differences in fruit and
flowering material cannot be matched with the
groups as based on the fruits.
The few specimens seen from New Ireland fit in
with group I. The populations from the Solomon
Islands have, as mentioned in the description, fruits
reaching larger sizes than the New Guinean plants
of group I, but with an ample overlap.
The shape, size, and consistency of the leaves
are somewhat variable in this species without,
however, permitting the distinction of infraspecific
taxa.
Some specimens with exceptionally long ra-
cemes and large leaves may resemble P. dolicho-
botrys, but differ in the hairy ovaries, fruits, and
seeds.
28. Prunus sclerophylla Kalkman, Blumea 13
(1965) 67. — Type: Robbins 573, Papua New
Guinea.
Small to medium-sized trees or shrubs. Twigs
densely hairy. Leaves elliptic to oblong, or ovat-
ish, 1.5-7 by 1-3 cm, base acute, apex acute or
shortly acuminate, coriaceous, with 4-7 pairs of
nerves, venation not conspicuous, hairy when
young, glabrescent but remaining hairy under-
neath, basal glands 0-2, flat. Petiole 0.2-0.5 cm
long. Stipules narrowly triangular, 3-4 by 0.7-1
mm, free. Racemes solitary, in axils of extant or
fallen leaves, up to 2.5 cm long, peduncle up to
0.5 cm, rachis hairy, pedicels up to 2.5 mm long.
Hypanthium 2-2.5 mm high, densely hairy out-
side. Perianth segments subequal or at least not
regularly differentiated as sepals and petals, 10-12,
1-1.5 mm long, hairy. Stamens 18-28, filaments
up to 3 mm, glabrous, anthers 0.5 mm long. Ovary
glabrous or with few hairs especially near suture,
style up to 2.5 mm long. Fruits transversely ellip-
soid, 6-8 by 7-9.5 mm, exocarp glabrous, black
when ripe, endocarp glabrous inside. Seed with
glabrous testa.
Distribution - Papua New Guinea, seen from a
number of mountains in W and E Highlands Prov-
inces.
Habitat - Forest, also mossy forest, and sub-
alpine shrubland, altitudes 2100-3100 m.
Uses - The bark of larger trees is, like that of
other species of Prunus, used for native waist-belts
(Pullen 148, Mt Hagen area).
Note - The extremely limited value of recording
vernacular names is well illustrated by the three
different names for this species, given in the course
of two weeks by one or more informants speaking
the Minj language to one collector, R. Pullen
(5197 yurih, 5226 beindangan, 5329 bugl-bakl)!
29. Prunus spicata Kalkman, Blumea 13 (1965)
69. — Type: Clemens 40755, Sabah.
Trees up to 12(-25) m, or large shrubs, bark
dark purplish or brownish. Twigs densely hairy.
Leaves elliptic to oblong or lanceolate to ovate,
6-18 by 2.5-6.5 cm, base rounded, apex acute,
long tapering or acuminate, herbaceous, with 6-
12 pairs of nerves, venation inconspicuous to in-
visible, (rather) densely hairy when young, hairs
usually remaining underneath, basal glands 0-2
(-4), flat. Petiole 0.2-1 cm long. Stipules nar-
rowly triangular to oblong, 3-8 by 0.5-2.5 mm,
free, rather persistent, margin hairy, with glands.
Racemes solitary, axillary, 1— 10(— 17) cm long,
peduncle 0-0.5 cm, rachis hairy, pedicels 0-1
mm long. Flowers sometimes male. Hypanthium
1.5-3 mm high, densely to sparsely hairy outside.
Perianth segments subequal or at least not regular-
ly differentiated as sepals and petals, 8-10, 0.5-2
mm long, hairy. Stamens 15-30, filaments up to
4 mm, glabrous or sparsely hairy at base, anthers
0.3-0.5 mm long. Ovary densely hairy, style up
to 4.5 mm long, often partly hairy, pistillode in
male flowers minute. Fruits globular to transver-
sely ellipsoid, 9-14.5 by 9-14.5 mm, exocarp
hairy to almost glabrous, red when ripe (becoming
black?), endocarp glabrous inside. Seed with gla-
brous testa.
Distribution - Borneo, Philippines (Luzon, a
sterile specimen from Mindoro).
Kalkman — Rosaceae
347
Habitat - Primary (and secondary?) forest, also
on riverbanks, at 1 100-1500 m altitude.
Note - Judging from the c. 8 fruiting specimens
seen, the fruits in Luzon are more globular, those
in Borneo more transversely ellipsoid and average-
ly somewhat smaller.
30. Prunus subglabra (Merr.) Kalkman, Blu-
mea 13 (1965) 87. — Pygeum subglabrum
Merr., Philipp. J. Sc. 30 (1926) 395. — Type:
Ramos & Edaho BS 45014, Mt Pulog.
Small trees up to 7 m. Twigs sparsely hairy
when young. Leaves elliptic, 6-10 by 4-5 cm,
base acute, apex shortly acuminate, coriaceous,
with 6-8 pairs of nerves, venation inconspicu-
ous, sparsely hairy when young, glabrescent, basal
glands 0-2, flat or slightly hollowed but not bulg-
ing above. Petiole 0.5-1.5 cm long. Stipules
elliptic, c. 8 by 3-4 mm, free, keeled inside,
sparsely hairy outside. Racemes solitary, in axils
of extant or fallen leaves, 2-5 cm long (in fruit-
ing stage), rachis stout, hairy, fruit pedicels up to
3 mm long. Flowers only known from some old
fragments. Hypanthium c. 3 mm high, densely
hairy outside. Perianth segments subequal, c. 9(?),
c. 2 mm long. Stamens c. 45, filaments up to 3.5
mm, anthers c. 0.5 mm long. Ovary hairy, prob-
ably densely so. Fruits subglobular, 15-20 by
(? 12-) 17- 19 mm, exocarp with few hairs, red,
endocarp sparsely hairy inside. Seed with glabrous
testa.
Distribution - Luzon: only seen from Mt Pulog
and nearby Mt Tabayog.
Habitat - Forest, one collection from mossy
forest, altitude 2400-2700 m.
Note - Two recent collections are both in fruit
and do not add to the incomplete information on
the flowers of this still badly known species, of
which only three collections seem to exist, al-
though Mt Pulog on Luzon is a well-explored
place.
3L Prunus turfosa Kalkman, Blumea 13 (1965)
90. — Type: Anderson 13123, Sarawak.
Small trees up to 12 m, rarely buttressed, bark
grey, smooth or slightly rough. Twigs sparsely
hairy or glabrous. Leaves elliptic to oblong, 8-
13(— 18) by 4-6.5 cm, base acute, rarely rounded,
apex obtuse to acuminate, herbaceous, with 8-12
pairs of nerves, venation inconspicuous, sparsely
hairy to entirely glabrous, basal glands 2, deeply
hollowed, in the contracted leaf-base. Petiole 0.5-1
cm long. Stipules not seen. Racemes in fascicles
of up to 4, in axils of extant or fallen leaves, up to
3.5 cm long, peduncle less than 0.5 cm, rachis
hairy, pedicels 1.5-2 mm long. Hypanthium 1.7-
2.5 mm high, sparsely hairy outside. Perianth seg-
ments subequal, 5-8, up to 1 mm long. Stamens
15-30, filaments up to 3.5 mm, glabrous or
slightly hairy at base, anthers 0.2-0.5 mm long.
Ovary sparsely hairy (to entirely glabrous?), style
up to 4.5 mm long. Fruits subglobular, 7-9 by
8-9 mm, exocarp with some hairs or glabrous,
colour unknown, endocarp glabrous inside. Seed
with glabrous testa.
Distribution - Borneo: Sarawak, Kalimantan
(one collection from W Kalimantan, near the bor-
der with Sarawak), Brunei [according to Anderson,
Check-list Trees Sarawak (1980) 295, but speci-
mens not seen].
Habitat - Peat swamp forest, about sea-level.
Note - New data about this species are not
available and it seems that mere has not been done
any collecting in the habitat of Prunus turfosa after
1961.
32. Prunus turneriana (F.M. Bailey) Kalk-
man, Blumea 13 (1965) 81. — Pygeum turner -
ianum F.M. Bailey, Bot. Bull. Queensl. Dep.
Agr. 8 (1893) 75; Queensl. Fl. 2 (1900) 525,
pi. 19. — Type: Cowley s.n., Queensland.
Prunus glomerata (Koehne) Kalkman, Blumea 13
(1965) 81. — Pygeum glomeratum Koehne, Bot
Jahrb. 52 (1915) 340. — Type: Ledermann 9497,
Papua New Guinea.
Trees up to 30 m, sometimes slightly buttressed,
bark smooth or with lenticels in longitudinal
fines, brown to gTey. Twigs hairy, glabrescent
Leaves elliptic to ovate or obovate, 7-23 by 4-1 1
cm, base rounded to acute, apex acute to rounded,
coriaceous to herbaceous, with 7— 12(— 15) pairs of
nerves, venation more or less transverse, usually
conspicuous beneath when dry, densely to sparsely
hairy when young, glabrescent, basal glands 2-6,
flat. Petiole 0.5-1.5 cm long. Stipules ovate to
triangular, sometimes large, 4— 7(— 15) by 1— 3(— 10)
mm, free. Racemes solitary, in axils of extant or
fallen leaves, 2-9 cm long, peduncle absent or
very short, rachis hairy, pedicels 0-2 mm long.
Flowers often male. Hypanthium (2-)3-4 mm
high, hairy outside and also inside. Perianth sub-
equal to more or less regular, 6-14, up to 1.5 mm
long. Stamens 15-50, filaments up to 4.5 mm,
often hairy at base, anthers 0.5-1.2 mm long.
Ovary hairy, style up to 5 mm long, hairy, pistil-
lode in male flowers minute. Fruits compressed
subglobular, 17-33 by 18-34 mm, hairy, black
348
Flora Malesiana ser. I, Vol. 11 (2) (1993)
when ripe, mesocarp rather thick when living, en-
docarp c. 1 mm thick, woody, usually hairy inside.
Seed with usually sparsely hairy testa, hairs often
especially near apex, sometimes glabrous, rarely
densely hairy.
Distribution - Moluccas (one specimen seen
from Bacan), Papua New Guinea (and one sterile
collection from Irian Jaya), Bismarck Archipelago
(only one specimen, from New Hanover), Australia
(N Queensland).
Habitat - Forest, from sea-level to 2400 m alti-
tude.
Notes - The species is characterized by its thick-
walled, compressed subglobular fruits. In Australia
the species is much more uniform than in New
Guinea, but on the evidence available now, it is
not possible to discriminate two taxa, as done in
the 1965 revision. In New Guinea the species is
found at low altitudes (as in Queensland) but also
in montane forest. In the higher altitudes the
leaves are often relatively small and densely hairy,
but the variation in these two characters is contin-
uous.
The fruits are very variable in dimension, also
within one specimen; the extremely large fruits all
come from New Guinea. The indumentum on the
seedcoat is in Australian specimens usually sparse
and often restricted to the apex. In New Guinean
specimens the seeds are sometimes quite glabrous,
rarely densely hairy.
33. Prunus versteeghii Kalkman, Blumea 13
(1965) 104. — Type: Versteegh BW4843, Irian
Jaya.
Trees up to 25 m, buttresses sometimes pres-
ent, bark grey(-brown), strongly smelling. Twigs
sparsely hairy to glabrous. Leaves oblong to ob-
long-ovate, 10-15 by 4-8 cm, base rounded to
acute, apex acute, herbaceous, with 7-9 pairs of
nerves, venation not conspicuous, glabrous to very
sparsely hairy, basal glands 2, flat. Petiole 0.7-1
cm long. Stipules (few seen) c. 6 by 2 mm, free.
Racemes solitary or in fascicles of 2-3, in axils
of extant or fallen leaves, 3-10 cm long, peduncle
very short, rachis hairy, pedicels 1-2 mm long.
Only male flowers seen. Hypanthium c. 2 mm high,
hairy outside. Sepals triangular, c. 1 mm long,
hairy. Petals elliptic to obovate, c. 1 mm long,
hairy. Stamens 35-45, filaments 1.5 mm, anthers
0.5-0.7 mm long. Pistillode in male flowers
minute, ovary in bisexual flowers probably hairy.
Fruits transversely ellipsoid to didymous, 15-19
by 22-28(-30) mm, exocarp sparsely hairy to
glabrous, purplish black when ripe, mesocarp
rather thick, endocarp glabrous inside. Seed with
glabrous testa.
Distribution - New Guinea, seen from five lo-
calities far apart
Habitat - Forest, probably preferring wet places.
Altitude 0-300 m.
34. Prunus wallaceana Kalkman, Blumea 13
(1965) 86. — Type: Kostermans 18587, Sum-
bawa.
Trees up to 30 m, rarely buttressed, bark smooth,
grey to darkbrown, with lenticels. Twigs hairy to
glabrous. Leaves elliptic to oblong, sometimes
ovatish, 10-18 by 4-8 cm, base usually rounded,
rarely more acute, apex gradually tapering to
shortly acuminate, herbaceous, with 8-14 pairs of
nerves, venation widely transverse, not conspicu-
ous, usually glabrous, rarely with some pubescence,
basal glands (0-)2, flat. Petiole (0.5-)l-2.5 cm
long. Stipules oblong, 3-9 by 0.8-2.3 mm,
free, sometimes slightly keeled inside, sometimes
with marginal glands. Racemes solitary, in axils
of extant or fallen leaves, 4-10 cm long, peduncle
very short, rachis sparsely hairy, pedicels (0.5 -)
3-6 mm long. Hypanthium 2-3 mm high, (spar-
sely) hairy outside. Perianth 5- or 6-merous. Sepals
triangular, 1—1.5 mm long, hairy outside. Petals
obovate, 1.5-3 mm long, hairy outside, white.
Stamens 35-55, filaments up to 4.5 mm, gla-
brous or hairy at base, anthers 0.3-0.7 mm long.
Ovary glabrous, style up to 6 mm long. Fruits
transversely ellipsoid, 10-13 by 13-18 mm, exo-
carp glabrous, red when ripe (or ultimately becom-
ing black?), endocarp glabrous inside. Seed with
glabrous testa.
Distribution - Celebes (S peninsula, Talaud I.),
Lesser Sunda Islands (seen from Sumbawa, Sumba,
Flores), Moluccas (seen from Ternate, Ceram).
Habitat - Primary and secondary forest, also
along rivers, altitude 0-1700 m.
Subgenus Padus
See under Insufficiently known species, Prunus C.
Kalkman — Rosaceae
349
Subgenus Amygdalus
In Malesia only cultivated
Prunus persica (L.) Batsch, Beitr. Entw. Pragm.
Gesch. 1 (1801) 30. — Amygdalus persica L.,
Sp. PI. (1753) 472. — Persica vulgaris Miller,
Gard. Diet (1768) 465. — Type: in LINN.
Deciduous shrubs or trees, up to 8 m. Twigs
glabrous. Leaves lanceolate, 5 — 8(— 15) by 1.5-2
cm, margin serrate, basal glands on top of petiole
or at base of blade in the margin. Flowers appear-
ing before the leaves, often in pairs next to axil-
lary buds, 5-merous, pedicels 0. Petals pink. Sta-
mens 35-40. Ovary hairy to glabrous. Fruits
globular, faintly furrowed longitudinally, 3-8 cm
diam., exocarp hairy (peach) or glabrous (nectarine),
yellow to red, mesocarp fleshy, yellow to red, en-
docarp thick and hard, deeply pitted and furrowed.
Distribution - Native in China, cultivation
spread from there, now cultivated wherever the
climate is suitable. In Malesia rarely cultivated in
montane areas, among others in East Java.
Uses - Fruits edible and according to many one
of the world's most delicious table fruits. See in-
formation on aspects of cultivation in tropical Asia:
S. Subhadrabandhu in E.W.M. Verheij & R.E.
Coronel (eds.), Edible fruits and nuts, Plant Res.
SE Asia (PROSE A Handbook) 2 (1991) 262-266.
Common names - Peach, nectarine (English),
Persik (Indonesia, Malaysia), Peras (Philippines).
INSUFFICIENTLY KNOWN SPECIES
Prunus odorata (Henderson) Whitmore in Tree
Fl. Malaya 2 (1973) 338. — Pygeum odoratum
Henderson, Gard. Bull. Str. Settl. 7 (1933) 101,
pi. 20. — Type: Henderson SFN 23278, Malaya.
Small trees. Twigs glabrous. Leaves ovate-
elliptic, 6.5-11 by 3.5-5.5 cm, base rounded to
acute, apex acute to acuminate, herbaceous, with
6-9 pairs of nerves, venation inconspicuous, gla-
brous, basal glands 2-4, flat. Petiole 0.5-1 cm
long. Stipules 4 by 1 mm, free. Racemes solitary,
up to 4 cm long, rachis hairy, pedicels up to 5 mm
long. Flowers fragrant (Henderson SFN 23278).
Hypanthium c. 2.5 mm high, hairy outside. Peri-
anth segments subequal, 7-9, 1-1.2 mm long.
Stamens 28-38, filaments up to 5 mm long.
Ovary hairy, style up to 5 mm long, hairy at base.
Fruits not seen.
Distribution - Malaya, Pahang, Cameron High-
lands.
Habitat - Nothing known except the altitude:
1440 m.
Note - Although now formally transferred from
Pygeum to Prunus, this taxon - if it is one - has
not become better known than in 1965, when I put
it under the 'incompletely known species', only
flowers and flowerbuds being present on the two
collections. There is a distinct likeness to Prunus
grisea var. tomentosa.
Prunus A
Trees up to 25 m high and 30 cm diam., bark
light to darkbrown, smooth. Twigs practically gla-
brous. Leaves elliptic, 14-17 by 8-10 cm, base
rounded to truncate, rarely more acute, apex shortly
acuminate to rounded, herbaceous, with 8-10 pairs
of nerves, venation transverse, not very conspicu-
ous, glabrous above, sparsely short-hairy below,
basal glands 2, deeply hollowed and bulging above,
situated in a contraction of the leaf-basae. Petiole
1-1.5 cm long. Stipules ovate, sometimes oblique,
6-9 by 1.5-3 mm, free(?), with glands on mar-
gin and (often?) with 1-3 flat to slightly hollowed
glands on the outer surface, sparsely hairy outside.
Inflorescence a compound raceme, in axils of
leaves or scars, up to 10 cm long, with up to 4
side-branches of 4-6 cm long, rachis sparsely
short-hairy, pedicels up to 1 mm long. Flowers
sometimes male. Hypanthium 1.5-2 mm high,
hairy outside. Perianth segments 8-10, subequal,
0.5-1 by 0.2-0.5 mm, hairy outside. Stamens
20-25, filaments up to 3.5 mm, glabrous or with
few hairs at base, anthers 0.2 mm long. Ovary gla-
brous, style up to 4 mm long, pistillode in male
flowers small. Fruits (only young ones seen) prob-
ably globular or transversely ellipsoid, 8 by 10 mm
when immature, glabrous. Seed with glabrous testa
(when young).
350
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Distribution - Borneo, seen from Sarawak and
Sabah.
Habitat - Forest on hillsides and riverbanks, up
to 450 m altitude.
Note - The leaves of the six collections, here
indicated as PrunusA (belonging to sectMeso-
pygeum), are of the same kind as those of Prunus
polystachya, having the large marsupial glands in
a contraction of the leaf -base.
The young fruits present on three of the speci-
mens do certainly not give the impression that they
could grow into the large fruits as in malayana or
polystachya.
Prunus B — Pygeum macropetalum Koehne,
Bot. Jahrb. 51 (1913) 198, 222; Baker f., J. BoL
62 (1924) Suppl. 33. — Types: Forbes 2343,
2354a, Sumatra, Mt Dempo.
Trees. Twigs glabrous. Leaves ovate-lanceolate
to elliptic or oblong, 8-17 by 3.5-9 cm, base
acute to rounded, apex tapering to acuminate, coria-
ceous, with 10-15 pairs of nerves, venation trans-
verse, not conspicuous, both surfaces glabrous
when mature, sparsely hairy below when very
young only, basal glands 0-2, flat. Petiole 1-2
cm long. Stipules 7-8.5 by 2-3 mm, free, gla-
brous, margin glandular. Racemes solitary, in the
axils of fallen leaves, 7-15 cm long, peduncle 1-2
cm long, rachis hairy, pedicels 4-8 mm long, pink.
Hypanthium 3-4 mm high, short-hairy outside.
Perianth differentiated as sepals and petals, but not
rarely with irregularities, 4- or 5-merous. Sepals
triangular, 1.5-2 mm long, hairy outside. Petals
suborbicular to elliptic, 2-3.5 mm long, hairy
outside, white. Stamens 35-55, filaments up to 7
mm, glabrous or with some hairs at base, anthers
0.7-1 mm long. Ovary glabrous, style up to 5 mm
long. Fruits subglobose-ovoid, 10-12 mm long,
see note. Seeds with glabrous testa.
Distribution - Sumatra, see note. Part of Forbes
2354a has 'Java' printed on the labels.
Habitat - Altitude 1700-2000 m.
Note - The two specimens cited with the origi-
nal description seem to be conspecific with De
Wilde & De Wilde-Duyfjes 14087 from Mt Ke-
tambe and with the sterile specimen Meijer 6434
from Mt Kerinci, both also from Sumatra but
rather wide apart. The species belongs to sect.
Mesopygeum and is obviously related to Prunus
grisea and P. ceylanica, but the very long racemes
and the rather large flowers are differentiating. It is
entered here because of the insufficient data on the
fruits. The description given rests on the field note
of De Wilde c.s. 14087, but on the herbarium
specimens only some immature fruits are present
and a possibly full-grown one which is subglobu-
lar and measures 8 by 7 mm, with some few hairs.
Prunus C — Prunus luzoniensis Merr. & Quis.,
nomen on herbarium sheet
Small trees. Twigs glabrous. Leaves lanceolate,
8-10.5 by 2.5-3 cm, base acute, margin finely
serrate, apex gradually narrowed, herbaceous, with
c. 15 pairs of nerves, between two primary nerves
often with an ± equally strong and parallel nerve,
venation not conspicuous, upper surface glabrous,
lower surface with hairs on and near midrib, doma-
tia absent, basal glands usually 2, on the top of
the petiole. Petiole 1-1.5 cm long. Stipules nar-
rowly triangular, 9-10 by 1-1.5 mm, free, on the
twig, both sides hairy, margin dentate-glandular.
Racemes terminal on lateral branches, bearing two
normal but small (up to 7.5 cm long) leaves in the
part below the flowers, entire branch 9-13 cm
long, with c. 20 flowers, rachis hairy, pedicels 2-3
mm long. Hypanthium c. 2.5 mm high, sparsely
hairy outside. Sepals 5, triangular with rounded
apex, c. 1.5 by 1.5-2 mm, margin with some
glandular teeth, (almost) glabrous outside. Petals
5, broadly obovate, 3-4 by 3.5-4.5 mm, gla-
brous, white. Stamens c. 20, filaments up to 3
mm, glabrous, anthers 0.7-1 mm long. Ovary
glabrous, style up to 1.5 mm long. Fruits not seen.
Distribution - One specimen seen from Mt
Pulog, Luzon, Philippines.
Habitat - The specimen comes from a "partly
open field, close to stream", 2000 m altitude.
Note - Celestino c.s. PNH 4337 is the first
and up till now the only collection of a Malesian
Prunus belonging to the subgenus Padus which is
characterized by the leaves on the basis of the ra-
ceme. It was collected in 1948 on a well-visited
mountain and it can be doubted whether it came
from a wild tree. Identification is difficult since too
many badly demarcated species have been described
in this subgenus. Also because of the absence of
fruits it seems better to keep it under the Insuffici-
ently known species than to create another syno-
nym which later will have to be reduced.
Prunus D
Tall trees. Twigs hairy. Leaves elliptic to ovate-
oblong, 11-16 by 4.5-7.5 cm, base acute, apex
gradually narrowing, coriaceous, with 7-9 pairs of
nerves, venation inconspicuous, glabrous above,
sparsely hairy below, basal glands 2, flat, at the
very base of the midrib. Petiole c. 1 cm long.
Kalkman
Rosaceae
351
Stipules not seen. Racemes solitary, only seen in
fruit, on old twigs, 1-2.5 cm long, stout, pedun-
cle almost absent, rachis hairy, fruiting pedicels up
to 2 mm long. Flowers only seen as remnants.
llypanlhium c. 2 mm high, densely hairy outside.
Perianth segments subequal, 10-12, 2-2.5 mm
long, hairy. Stamens short, filaments up to 2 mm,
anthers 0.6 mm long. Fruits compressed (broadly)
ovoid, 16-18 by 14-18 mm, exocarp sparsely
hairy, especially at top, green when ripe (?), meso-
carp thin, endocarp thin (c. 0.5 mm), sparsely hairy
inside. Seed with glabrous testa.
Distribution - One specimen seen (Hoogland &
Craven 11086), from E Sepik Province, Papua New
Guinea.
Habitat - Rain forest, at 1 160 m altitude.
Note - This species belongs to sect. Meso-
pygeum and seems to be rather well distinct. The
fruits are not like any described species, much too
large for Prunus grisea, too small for P. turneri-
ana, two species which could be considered.
EXCLUDED
Prunus zippeliana Miq., Fl. Ind. BaL I, 1
(1855) 367. — Type: Zippel s.n., Java, holo
in L, sheet 908.202-880.
Obviously collected from a cultivated tree, prob-
ably in the Botanic Garden at Buitenzorg (Bogor)
and originating from Japan. The natural range of
the species is China, Taiwan, Japan, and Vietnam.
Several Pygeum species names cannot be account-
ed for, types being lost or insufficient. See lists of
'Incompletely known species' and 'Dubious spe-
cies' in Kalkman (1965).
Flora Malesiana ser. I, Vol. 1 1 (2) (1993) 353-373
AMARYLLIDACEAE
(including Hypoxidaceae)
(D.J.L. Geerinck, Brussels, Belgium)
Perennial herbs with bulbs, tubers or rhizomes. Leaves simple, with parallel nerves. In-
florescences terminal or axillary, in cymes, spikes or umbels (in Amaryllidoideae), or
flowers solitary, bracteate and often with one or few spathes (in Amaryllidoideae). Flowers
bisexual, actinomorphic or zygomorphic, sometimes marcescent. Tepals in 2 whorls, free
or united into a tube, sometimes with a conspicuous corona. Stamens 6, free or some-
times united into a false corona, often inserted at the mouth of the perigone-tube; anthers
basifixed, dorsifixed or medifixed, often versatile. Ovary inferior, 3-celled with axillary
placentas; ovules 1 to numerous per cell. Fruit capsular, dehiscing either loculicidally or
irregularly, or fruit a berry. Seeds globose or flattened, sometimes winged.
Distribution — Cosmopolitan, with c. 80 genera and around 1000 species. In Malesia
only 6 genera are indigenous or naturalized, but many others are cultivated in botanic and
private gardens (see the list on p. 371).
Taxonomy — The family is treated here in a broad sense, comprising the genera with
an inferior ovary, i.e. excluding the Allioideae (= Alliaceae), which are characterized by a
superior ovary. In Malesia there are no indigenous species of the latter family, which is
treated elsewhere in this instalment (p. 375).
The Agavoideae (partly with an inferior ovary and partly with a superior one) are also
excluded. The family Agavaceae has one indigenous genus in Malesia (Dracaena, includ-
ing Pleomele).
In the Amaryllidaceae two subfamilies are here recognized which are often considered
to be distinct families: the Amaryllidoideae (= Amaryllidaceae s.str.), with umbellate in-
florescences and spathes, and the Hypoxidoideae (= Hypoxidaceae) which are never um-
bellate and are without spathes.
Haemodoraceae were treated by Van Steenis in Flora Malesiana I, 5 (1955) 1 1 1-113
[and see Additions in Fl. Males. I, 10 (1989) 717].
Palynology — Pollen of Amaryllidaceae has been studied in detail by Radulescu
(1972), Schulze (1983), Meerow (1987, 1989), and Meerow & Dehgan (1988). Amaryl-
lidaceae pollen grains are small to very large (largest equatorial axis 14-158 urn). Small
grains (usually < 30 um) are found in the Gale .theae (Galanthus, Leucojum p.p.). Very
large pollen (> 100 um) occurs only in Hymenocallis p.p. Tetragonal tetrads are known
in Stenomesson elwesii (Meerow et al. 1986).
The apertural system is monosulcate or dicolpate. Sulci are nearly as long as the longest
equatorial axis, or extend slightly on the proximal grain side. The position of the colpi in
dicolpate pollen with respect to the distal and proximal poles is unknown.
(353)
354 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Exine stratification is generally distinct. Electron micrographs show a columellate, tec-
tate or semitectate sexine in monosulcate pollen. Zavada (1983) has demonstrated a very
thin, finely granular infratectum and a thick nexine traversed by minute channels in the
dicolpate pollen of Crinum americanum (Amaryllidaceae). An intectate, gemmate sexine
was found in the tetrads of Stenomesson elwesii.
Ornamentation is reticulate or sparsely scabrate-microechinate, rarely psilate/punctate
(Galantheae) to verrucate/fossulate (Eucrosia). The lumina in reticulate patterns are small
to very large (up to c. 10 urn in Hymenocallis). Usually lumen size decreases towards the
aperture, and often it is distinctly less at the apocolpial ends of a grain. In several Hymeno-
callis species these parts ('auriculae') have an ornamentation quite different and clearly
separated from that of the main part of the grain (Ravikumar & Nair 1982; Meerow &
Dehgan 1985).
Two main pollen types may be distinguished in the family: 1) monosulcate reticulate
pollen, and 2) dicolpate scabrate-microechinate pollen. Infratectum structure might be
another important feature separating both types (see above). The latter characterizes all
genera of the Amaryllideae (sensu Dahlgren et al. 1985; see also Schulze 1983 and Snij-
man 1991), and Pauridia of the Hypoxidoideae (Thompson 1979; Simpson 1983). The
former type is found in all other Amaryllidaceae. The two colpi in Amaryllideae pollen are
on opposite grain sides, whereas in Pauridia pollen they are situated in the same hemisphere.
Also the nature of the scabrate ornamentation is different in Pauridia. Other Hypoxidoi-
deae have monosulcate, finely reticulate pollen, more or less the same as in most Amaryl-
lidoideae and many other monocots.
References: Dahlgren, R.M.T., H.T. Clifford & P.F. Yeo, The families of the Monocotyledons
(1985) 199. — Meerow, A.W., Syst. Bot. 12 (1987) 460-492; Ann. Missouri Bot. Gard. 76 (1989)
136-220. — Meerow, A.W. & B. Dehgan, Amer. J. Bot. (1985) 540-547; Amer. J. Bot. 75 (1988)
1857-1870. — Meerow, A.W., N.B. Dehgan & B. Dehgan, Amer. J. Bot. 73 (1986) 1642-1644. —
Radulescu, D., Acta Bot. Hort. Bucur. 1970-71 (1972) 245-273. — Ravikumar, C. & P.K.K. Nair,
J. Palynol. 18 (1982) 43-54. — Schulze, W., Wiss. Z. Fr.-Schiller-Univ. Jena, Math.-Naturwiss. R. 32
(1983) 985-1003. — Simpson, M.G., Grana 22 (1983) 79-103. — Snijman, D.A., Bothalia 21 (1991)
125-128. — Thompson, M.F., Bothalia 12 (1979) 621-625. — Zavada, M.S., Bot. Review 49 (1983)
R.W. J.M. van der Ham
Phytochemistry — Difficulties in demarcating families in Liliales are aptly discussed
by Cronquist (1981, 1988). He accepted Iridaceae, Haemodoraceae, Alo(e)aceae, Agava-
ceae, Xanthorrhoeaceae and Smilacaceae as distinct families, but united Amaryllidaceae
s. str. and all modern segregates from Liliaceae and Amaryllidaceae sensu Pax et Hoffmann
(e.g. Agapanthaceae, Alliaceae, Alstroemeriaceae, Asparagaceae, Asteliaceae, Antherica-
ceae, Aphyllanthaceae, Calochortaceae, Colchicaceae, Convallariaceae, Dianellaceae,
Hemerocallidaceae, Hyacinthaceae, Hypoxidaceae, Herreriaceae, Melianthiaceae, Rusca-
ceae, Tecophilaeaceae, Tricyrtidaceae, Trilliaceae and others) in Liliaceae sensu Cronquist,
because there seem to be no characters available for an unequivocal demarcation of such
smaller families. In some instances phytochemistry may be able to assist taxonomists in
delimitating taxa of subfamily or family rank. This seems exactly to be the case with
Alliaceae, Amaryllidaceae s.str., Hypoxidaceae and Haemodoraceae [the latter already
Geerinck — Amaryllidaceae 355
treated in Flora Malesiana I, 5 (1954) 1 1 1-1 15, but without a paragraph on phytochem-
istry]. Therefore the taxonomically most promising secondary metabolites of the four taxa
mentioned will be discussed shortly sub Alliaceae and Amaryllidaceae. The chemistry and
chemotaxonomy of all four taxa was treated more comprehensively by Hegnauer (1963,
1986). Chelidonic acid seems to be a major organic acid in all of them and in most Lilia-
ceae sensu Cronquist.
Amaryllidaceae s.str. (= Amaryllidoideae sensu Melchior 1964). — Alkaloid produc-
tion and storage is a key character of this taxon (Hegnauer 1990). Amaryllidaceous or lyco-
rine-type alkaloids seem to be present in every species of Amaryllidoideae and have never
been found outside this taxon. Lycorine-type alkaloids represent a biosynthetically dis-
tinct type of alkaloids. They originate by condensation of a hydroxy benzaldehyde (C6-C1-
moiety) derived from phenylalanine with tyramine (C6-C2-moiety) derived from tyrosine.
The resulting dicyclic belladine-type bases are intermediates on the way to the more
numerous tri- and tetracyclic amaryllidaceous alkaloids. These alkaloids are toxic and are
stored often in combined forms such as glycosides, as glycosides acylated by fatty acids
and as phospholipids (Ghosal et al. 1985, 1991). Moreover, oxidation to quaternary com-
pounds (e. g. anhydrolycorinium, ungeremine) which may be subsequently transformed
to Schiff bases [Ri,R.2 = C = N-R3 like crinasiatine (Ghosal et al. 1988)] occurs under
certain conditions. Ghosal et al. (1990, 1991) described an astonishingly perfect co-evo-
lution between the specialised moth Polytela gloriosa and its host plants Amaryllis bella-
donna and vittata (= Hippeastrum vittatum), Crinum asiaticum and latifolium and Pan-
cratium biflorum. The moth makes use of toxic amaryllidaceous alkaloids for defensive
purposes and transforms ingested bases for storage in its own glycosides and acylated
glycosides, and the plant responds to the stress of herbivory by releasing free, more toxic
alkaloids from stored combined forms in young fruits. Moreover, both the moth and
several of its host plants respond to certain stress conditions with synthesis of one and the
same alkaloidal conjugate termed telastaside (Ghosal et al. 1991). Kumar & Roy (1985)
recorded a strong allelopathic effect for lycorine; this alkaloid inhibits germination of fern
spores and kills prothallia.
Phenolic compounds are represented by several classes. Caffeic acid and glycosides of
the flavonols kaempferol and quercetin seem to be nearly ubiquitous. Acetophenone deri-
vatives were isolated from species of B up hane (acetovanillone), Lycoris (trimethyl ether
of phloracetophenone) and from Pancratium biflorum [several derivatives of phloraceto-
phenone (Ghosal et al. 1989)]. Oddly substituted flavans, flavan-3-ols, chalcones and
2-methylchromones were reported for Hippeastrum ananuca. Narcissus pseudonarcissus
and isolated from bulbs and flowering stalks of Pancratium biflorum (Ghosal et al. 1989).
Chromones and flavans were reported also for bulbs of Pancratium maritimum (Ali et al.
1990) and flavans and chalcones are present in bulbs of Crinum augustum (Abd El-Hafiz
et al. 1990). Obviously these phenolic compounds are phytoalexins in some instances
{Narcissus pseudonarcissus). A strange type of parasitism was described by Ghosal et al.
(1986). Stolons of the grass Imperata cylindrica penetrate bulbs of Pancratium biflorum,
cause cell necrosis and initiate synthesis of red coloured stress compounds (flavans, chal-
cones) and release of cytotoxic free alkaloids from their conjugates. Piscidic acid is known
from Narcissus poeticus. and Lycoris radiata yielded a deoxypiscidic acid. These two aro-
356 Flora Malesiana ser. I, Vol. 11 (2) (1993)
matic acids may be connected with tyrosine metabolism and may have /?-hydroxyphenyl-
pyruvic acid and acetic acid or an equivalent C2 compound as building stones.
Saponins seem to be totally lacking in true Amaryllidaceae.
Amaryllidaceae store starch in vegetative storage organs. In many taxa starch is accom-
panied by water-soluble carbohydrates, fructans or glucomannans. The latter may become
the main carbohydrate reserve in a number of taxa. A tendency to replace starch by water-
soluble carbohydrates in vegetative storage organs of perennial plants is perceptible in many
other more or less cosmopolitan families of monocotyledons, e.g. Agavaceae, Gramineae,
Iridaceae, Liliaceae s.l.
Hypoxidaceae (Hypoxidoideae) — Glycosidic 1,5- and 1,3-diarylpentanes seem to be
key metabolites of Hypoxidaceae, which comprise many species of importance to tradi-
tional medicine in Africa and Asia including China and Japan. Rooperol, an 1,5-diaryl-
penta-l-en-4-yn with four phenolic hydroxy Is is the aglycone of hypoxoside and the obtu-
sosides of Hypoxis nyasica, obtusa, rooperi and others. Nyasol, l,3-diaryl-l,4-pentadien
with two phenolic hydroxyls is the aglycone of several glycosides from H. angustifolia
(Sibanda et al. 1990) and nyasicol, a l,5-diaryl-l,2-dihydroxypenta-4-yn with four phe-
nolic OH's, is the aglycone of several glycosides of H. nyasica (Galeffi et al. 1987), H.
angustifolia, interjecta and multiceps (Sibanda et al. 1990; Marini-Bettolo et al. 1991).
Nyasicoside was also isolated from rhizomes of an African species cited zsCurculigo re-
curvata (Chifundera et al. 1991). Biosynthetically these C6-C5-C6 phenolic aglyca are prob-
ably related to lignans which arise by condensation of two phenylpropane (C6-C3) build-
ing stones and have a C6-C3 + C3-C6 skeleton. Because one C-atom is lost, such C6-C5-
C6 compounds are also termed norlignans, norneolignans or even nonlignans or conioids;
they were formerly studied intensively in conifers (see Hegnauer 1986: 479, 481, 496).
Acuminatoside from rhizomes of Hypoxis acuminata (Bredenkamp et al. 1989) and
whole plants of//, obtusa (Msonthi et al. 1990) is the apiosylglucoside of geraniol.
Flavonoids are represented in the family by glycosides of kaempferol and quercetin.
Rhizomes and tuberous roots of Curculigo orchioides yielded a derivative of 2,3-dihydro-
myricetin.
Simple phenolic glycosides seem to be rather common in Hypoxidoideae. From whole
plants of//, obtusa Msonthi et al. (1990) isolated also obtusaside, a glucoside of gentisyl
alcohol which has OH-6 of its glucose esterified by 2,6-dimethoxy-3-hydroxybenzoic
acid. Rhizomes of C. orchioides contain curculigoside, which is 2',6'-dimethoxysalirepo-
side, and corchioside-A, which is a xyloglucoside of orcinol (Garg et al. 1989).
Curculigo orchioides is a famous medicinal plant from India and China known in Ayur-
vedic medicine as 'Kali Musli' (Pandey et al. 1983) and in China as 'Xiao Mao' (Jun-Ping
Xu et al. 1992). The lipid fraction of this crude drug (rhizomes + roots) yielded interest-
ing alkanones, the cycloartane derivative curculigol (Misra et al. 1990), a new pentacyclic
triterpene (Mehta & Gawarikar 1991) and several phytosterols. Yun-Ping Xu et al. (1992)
isolated a saponin mixture containing curculigosaponins-A to -J, all with the same sapo-
genin, curculigenin, which is a cycloartane derivative. Obviously in contrast to many
Liliaceae s.l. (see e.g. sub Alliaceae), Dioscoreaceae and other families of monocotyle-
dons, Hypoxidoideae do not produce steroidal C27-sapogenins, but triterpenic sapogenins
with the tetracyclic skeleton of cycloartane.
Geerinck — Amaryllidaceae 357
Haemodoraceae — Yellow to red 9-phenylphenalenones and biogenetically related pig-
ments are chemical markers of true Haemodoraceae. They are deposited in subterranean
parts, leaf bases, flowers, fruits and seeds and occur free (e.g. haemodorin) and as glyco-
sides (e.g. haemocorin). Phenalenone pigments were isolated from species of Anigozan-
thos, Conostylis, Haemodorum, Lachnanthes, Macropidia, Phlebocarya, Wachendorfia
and Xiphidium. Quite recently a new phenalenone pigment, thyrsiflorin, was isolated
from the yellow flowers of Wachendorfia thyrsiflora, which contains two other phenale-
nones in its roots (Dora et al. 1990). Obviously the North American genus Lophiola does
not belong to Haemodoraceae; its two species do not produce phenalenone pigments.
Phenalenones also occur in some Fungi; fungal phenalenones have a polyketide origin,
whereas Haemodoraceae produce their pigments from one phenylalanine, one tyrosine
and one acetate.
References: Abd El-Hafiz, M. A., et al., J. Nat. Prod. 53 (1990) 1349. — Ali, A. A., et al., Photo-
chemistry 29 (1990) 625. — Bredenkamp, N.W., et al., Phytochemistry 28 (1989) 263. — Chifundera,
K., et al., Tetrahedron 47 (1991) 4369. — Cronquist, A., An integrated system of classification of flower-
ing plants, Columbia Univ. Press (1981); The evolution and classification of flowering plants, 2nd ed.,
New York Bol Garden (1988). — Dora, G, et al., Planta Medica 56 (1990) 569. — Galeffi, C, et al.,
Tetrahedron 43 (1987) 3519. — Garg, S.N., et al., Phytochemistry 28 (1989) 1771 (also alkanols, phyto-
sterols, cycloartenol). — Ghosal, S., et al., Phytochemistry 24 (1985) 1825 (glucoalkaloid kalbreclasine),
2141 (chemistry and biology of Crinum alkaloids: review), 2703 (acyloxy alkaloid palmilycorine and
acylglucosyloxy alkaloid lycoriside from Crinum asiaticum); ibid. 25 (1986) 1097; ibid. 27 (1988) 1849
(new types of alkaloids, e.g. craugsodine, isocraugsodine, crinasiatine from fruits of Crinum asiaticum);
ibid. 28 (1989) 3193; ibid. 29 (1990) 805; Indian J. Chem. 30B (1991) 260 (telastaside). — Hegnauer, R.,
Chemotaxonomie der Pflanzen 2 (1963); 7 (1986): Alliaceae 281, 315-325, 488-489, 501, 685-731,
803; Amaryllidaceae s.str. 53-70, 475, 573-580; Haemodoraceae 228-230, 657-660; Hypoxidaceae
234-236, 661-663. — Hegnauer, R., in P. Baas, R. Geesink & C. Kalkman, The plant diversity of
Malesia, Kluwer Acad. Publ., Dordrecht (1990) 97. — Kumar, G. & S.K. Roy, Indian J. Exp. Biol. 23
(1985) 356. — Marini-Bettolo, G.B., et al., Tetrahedron 47 (1991) 6717. — Mehta, B.K. & R. Gawarikar,
Indian J. Chem. 30B (1991) 986. — Melchior, H. in Engler, Syll. Pflanzenfam. 2 (1964) 528. — Misra,
T.N., et al., Phytochemistry 29 (1990) 929. — Msonthi, J.D., et al., Phytochemistry 29 (1990) 3977.
— Pandey, H.C., et al., Intern. J. Crude Drug Res. 21 (1983) 33. — Sibanda, S., et al., Biochem. Syst.
Ecol. 18 (1990) 481. — Xu, Jun-Ping, et al., Phytochemistry 31 (1992) 233; Planta Medica 58 (1992)
R. Hegnauer
KEY TO THE GENERA
(Genera of cultivated species included)
la. Inflorescence always an umbel with 1 to few spathes (Amaryilidoideae) 2
b. Inflorescence not an umbel, spathes absent (Hypoxidoideae) 18
2a. Filaments inserted on the throat of the perigone-tube, with a thin membrane some-
times 1- or 2-toothed between the filaments (false corona). True corona absent . . 3
b. Filaments free from the perigone-tube, false corona absent. True corona present or
not 6
3a. Leaves distinctly petiolate, subcircular to elliptic, cuneate to slightly cordate at base . 4
b. Leaves sessile or subpetiolate, linear to narrowly elliptic, attenuate into the petiole 5
358 Flora Malesiana ser. I, Vol. 11 (2) (1993)
4a. Perigone-tube up to 3.5 cm long, straight. Ovules 2 per cell . . Proiphys (p. 363)
b. Perigone-tube at least 3.5 cm long, often curved. Ovules usually 6 or more per cell
Eucharis (cult., p. 371)
5a. Ovary with 10 or more biseriate ovules in each cell. Fruit dry, 3-loculicid. Seeds
black Pancratium (p. 362)
b. Ovary with 1-9 ovules near the bottom in each cell. Fruit fleshy, finally rupturing
laterally. Seeds green Hymenocallis (p. 361)
6a. Flowers with a corona Narcissus (cult., p. 373)
b. Flowers without a corona 7
7a. Inflorescences always 1-flowered 8
b. Inflorescences 2- to many-flowered 10
8a. Perigone zygomorphic with free tepals; the outer ones much broader than the inner,
the latter connate into a channel enveloping stamens and style
Sprekelia (cult., p. 373)
b. Perigone actinomorphic with tepals united at the base 9
9a. Perigone-tube less than 3 cm long. Longest filaments more than 10 cm long. Style
bent out of the flower Zephyranthes (cult., p. 373)
b. Perigone-tube more than 5 cm long. Longest filaments less than 8 cm long. Style erect
in the centre of the flower Cooperia (cult., p. 371)
10a. Tepals free, brilliant white with a small green blotch below the top
Leucojum (cult., p. 372)
b. Tepals at base united into a tube 11
11a. Rowers distinctly zygomorphic 12
b. Flowers actinomorphic or nearly so 14
12a. Leaves petiolate, 2 or 3 times as long as wide. Perigone violet. Ovules 2 per cell
Griffinia (cult., p. 372)
b. Leaves sessile, more than 5 times as long as wide. Perigone red, reddish, pink or
white. Ovules numerous in each cell 13
13a. Peduncle solid Amaryllis (cult., p. 371)
b. Peduncle fistular Hippeastrum (cult., p. 372)
14a. Perigone-tube more than 5 cm long Crinum (p. 359)
b. Perigone-tube up to 5 cm long 15
15a. Peduncle fistular. Stamens inserted in the perigone-tube. Fruit a capsule 16
b. Peduncle solid. Stamens inserted at the throat or on the segments of the perigone.
Fruit a berry 17
16a. Perigone-tube funnel-shaped Cyrtanthus (cult., p. 371)
b. Perigone-tube narrow Vallota (cult., p. 373)
17a. Perigone-tube funnel-shaped, segments wide, orange-coloured
Clivia (cult., p. 371)
b. Perigone-tube narrow, segments linear, reddish to purple Scadoxus (cult., p. 373)
1 8a. Tepals very shortly to distinctiy united into a tube. Fruit fleshy, indehiscent or irreg-
ularly dehiscent, often persistentiy beaked Curculigo (p. 366)
b. Tepals free. Fruit dry, regularly dehiscent, never beaked .... Hypoxis (p. 370)
Geerinck — Amaryllidaceae 359
Subfamily Amaryllidoideae
Inflorescence umbellate, or flowers solitary, and always with one or few spathes. Flowers
actinomorphic or zygomorphic. Tepals free or united into a tube, sometimes with a con-
spicuous corona. Stamens free or sometimes united into a false corona.
The majority of the genera and species belong to this subfamily.
CRINUM
Crinum L., Sp. PI. (1753) 290. — Type species: Crinum americanum L.
Bulbous herbs. Leaves radical, rarely biseriate, sessile or subpetiolate. Inflorescences
umbellate, many- to 1 -flowered, with 2 spathes; scape lateral, solid. Flowers white or
pinkish, sometimes with a red keel. Tepals subequal, united at the base into a long and
narrow tube. Stamens inserted in the throat; filaments free; anthers dorsifixed. Ovary with
1 to many ovules per cell; style filiform; stigma entire. Fruits indehiscent or irregularly de-
hiscent, with a fleshy or membranous wall. Seeds globose or somewhat flattened, greenish
or greyish. — Fig. 1.
Distribution — About 100 species in the tropics; 2 indigenous species in Malesia. Other
species are cultivated in gardens and may escape sometimes.
KEY TO THE SPECIES
la. Leaves narrowly to broadly elliptic, wider than 3.5 cm. Anthers 12-35 mm long
1. C. asiaticum
b. Leaves oblong to narrowly elliptic, to 3.5 cm wide. Anthers 7-9 mm long
2. C. gracile
1. Crinum asiaticum L., Sp. PI. (1753) 292; Crinum sumatranum Roxb., Hort. Beng. (1814)
Ker-Gawl., Curt. Bot. Mag. 27 (1808) t. 1073; 23; Miq., Fl. Ned. Indie 3 (1859) 581; Lindley,
Miq., Fl. Ned. Indie 3 (1859) 580; Blanco, Fl. Edw. Bol Reg. 13 (1887) L 1049; Naves, Nov.
Filip. ed. 3, 1 (1877) 314, t. 168; Naves, Nov. App. (1880) 254; Baker, Handb. Amaryll. (1888)
App. (1880) 254; Baker, Handb. Amaryll. (1888) 75. — Type: Campbell, Sumatra.
75; Koord., Meded. Lands Plantent. 19 (1898) Crinum macrantherum Engler, BoL Jahrb. 7 (1886)
309; Ridley, Mat. Fl. Mai. Penins. 2 (1907) 448; Peekel, Fl. Bismarck Arch. Natur. (1984)
68; Hallier, Nova Guinea 8 (1913) 899; Laut., 87. — Type: Naumann 75, Bismarck Archi-
Bot. Jahrb. 50 (1913) 302; Ridley, Fl. Mai. pelago.
Penins. 4 (1924) 301; Merr., Enum. Philipp. Crinum nonhianum Baker, Gard. Chron. 1 (1882)
Flow. PI. 1 (1925) 210; Henderson, Mai. Wild 671; Handb. Amaryll. (1888) 82; Ridley,
Fl. (1954) 164; Backer & Bakh. f., Fl. Java 3 Agr. Bull. 3 (1904) 310; Fl. Mai. Penins. 4
(1968) 136 (s.l.). — Types: India, Malabar (1924) 301. —Type: North 424, Borneo, Sara-
Coast and Sri Lanka in LINN. wak.
Crinum amabile Donn, HorL Cantabr. ed. 6(1811) Crinum cortifolium Hallier, Nova Guinea 8 (1913)
83; Ker-Gawl., Curt. Bot. Mag. 39 (1814) t. 900. — Type: von Romer 189, New Guinea.
1605; Miq., Fl. Ned. Indie 3 (1859) 580; Naves, Crinum macrophyllum Hallier, Nova Guinea 8
Nov App. (1880) 254; Baker, Handb. Amaryll. (1913) 899. — Type: Versteeg 1219, New Gui-
(1888) 75. — Type: Hort. Kew. Sumatra nea.
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Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 1. Crinum asiaticum L. Habit of flowering plant. New Guinea, bank of Merau R., NE of Merauke,
Irian Jaya (Photograph P. van Royen, 1954).
Crinum rumphii Merr., Interpr. Rumph. Herb. Am-
boin. (1917) 141. — Type: Robinson PI.
Rumph. Amb. 131, Ambon.
Haemanthus pubescens auct. non L. f.: Blanco, Fl.
Filip. (1837) 253.
Crinum giganteum auct. non Andr.: Blanco, Fl.
Filip., ed. 2 (1845) 175.
Crinum defixum auct. non Ker-Gawl.: Ridley, Mat.
Fl. Mai. Penins. 2 (1907) 68; Fl. Mai. Penins.
4(1924)301.
Herb to 200 cm tall. Leaves narrowly to broad-
ly elliptic, 50-150 by (3-)3.5-20 cm. Umbels
10-50-flowered; peduncle 3.5-12.5 cm long;
spathes 9-16 cm long; pedicels 1-5 cm. Flowers
white, fragrant at night. Perigone-tube 7.5-13 cm;
lobes 6-12.5 by 4-12 mm. Stamens pink to pur-
ple; filaments 3.5-7 cm long; anthers straight,
12-35 mm long. Seeds 1-5 per cell. - Fig. 1.
Distribution - Continental Asia (India, Burma,
Thailand, Laos, Cambodia, Vietnam), Cocos Is.
(Indian Ocean), Mascarenes, NW Australia, Pacific
Is. (Guam, Carolines, Marshalls, Solomons, New
Caledonia, Fiji, Samoa); Malesia: Sumatra, Ma-
laya, Singapore, Borneo (Brunei, Sarawak), Java,
Madura, Philippines (Luzon, Mindoro, Palawan,
Mindanao), N Celebes, Timor, Moluccas (Ambon,
Seram), New Guinea (Merauke, Sepik, Milne Bay),
Bismarck Archipelago.
Habitat - Sandy places, also coastal; riverine
forests.
Uses - Leaves and bulbs have medicinal uses as
emetic, etc. in India (Quisumbing, Medic. PI. Phi-
lipp. 1951, 171).
Geerinck — Amaryllidaceae
361
2. Crinum gracile E. Meyer in Presl, Rel.
Haenk. 1 (1827) 120; Naves, Nov. App. (1880)
255; Baker, Handb. Amaryll. (1888) 81; Merr.,
Enum. Philipp. Flow. PI. 1 (1925) 210.—
Type: Haenke, Luzon.
Crinum cumingii Baker, Gard. Chron. 2 (1881) 72;
Handb. Amaryll. (1888) 81; Vidal, Phan. Cu-
ming. Philipp. (1885) 152; Rev. PI. Vase. Filip.
(1886) 275. — Type: Cuming 1382, Luzon.
Herb to 80 cm tall. Leaves oblong to narrowly
elliptic, 30-60 by 1-3.5 cm. Umbels 5-12-flow-
ered; peduncle 15-30 cm long; spathes 5-7 cm
long; pedicels 0.5-2 cm long. Flowers white.
Perigone-tube 5-12 cm long; lobes 5-8 by 5-10
mm. Stamens purple; filaments 5-7 cm long;
anthers slightly curved, 7-9 mm long. Style pur-
ple. Seeds unknown.
Distribution - Malesia: Sumatra (Palembang),
Philippines (Luzon, Mindoro, Mindanao), New
Guinea.
Habitat - Primary and savanna forests at low
altitudes.
CULTIVATED SPECIES
Crinum finibriatum Baker — Backer & Bakltf.,
Fl. Java 3 (1968) 137.
Native to tropical Africa; cultivated as an orna-
mental. Umbel 3-8-flowered; perigone whitish to
purplish.
Crinum jagus (Thomps.) Dandy — Crinum gi-
ganteum Andr.; Backer & Bakh. f., Fl. Java 3
(1968) 137.
Native to tropical Africa; cultivated as an orna-
mental. Umbel 2-12-flowered; flowers fragrant;
perigone white.
Crinum kirkii Baker — Backer & Bakh. f., Fl.
Java 3 (1968) 137.
Native to tropical Africa; cultivated as an orna-
mental. Umbe! 12- 20- flowered: pengone greenish.
Crinum latifolium L. — Merr., Enum. Philipp.
Row. PI. 1 (1925) 210; Steiner, Philipp. Or-
nam. PI. (1960) 137; Peekel, Fl. Bismarck Arch.
Natur. (1984) 87. — Crinum moluccanum
Roxb.; Miq., Fl. Ned. Indie 3 (1859) 581.
Native to tropical Asia; cultivated as an ornamen-
tal and perhaps escaped. Umbel 10-20-flowered;
flowers fragrant; perigone green tinged with red.
Crinum longiflorum Thunb. — Backer &
Bakh. f., Fl. Java 3 (1968) 136.
Native to South Africa; cultivated as an orna-
mental. Umbel 6-16-flowered; flowers fragrant;
perigone greenish.
Crinum moorei Hook f. — Backer & Bakh. f.,
Fl. Java 3 (1968) 136.
Native to South Africa; cultivated as an orna-
mental. Umbel 6-12-flowered; perigone pinkish.
Crinum x powellii Baker — Backer & Bakh. f.,
Fl. Java 3 (1968) 136.
Hybrid between the two preceding species; cul-
tivated as an ornamental.
Crinum pusillum auct. non Herbert?
Fl. Ned. Indie 3 (1859) 581.
Doubful species.
Miq.,
Crinum sanderianum Baker — Backer &
Bakh. f., Fl. Java 3 (1968) 136.
Native to tropical Africa; cultivated as an orna-
mental. Umbel 3-6-flowered; perigone white with
a pink top.
Crinum zeylanicum L. — Merr., Enum. Phi-
Upp. How. PI. 1 (1925) 211; Backer & Bakh. f.,
Fl. Java 3 (1968) 137.
Native to tropical Africa and Asia; cultivated as
an ornamental and subnaturalized. Umbel 6-25-
flowered; flowers fragrant; perigone white with
wide pink bands.
HYMENOCALLIS
Hymenocallis Salisb., Trans. Hort. Soc. Lond. 1 (1812) 338. — Type species: Hymeno-
callis littoralis (Jacq.) Salisb.
Bulbous herbs. Leaves radical, sessile or subpetiolate. Inflorescences umbellate, many-
to 2-flowered, with 2 spathes. Flowers white. Tepals equal, united into a long tube; seg-
ments erect to patent. Stamens inserted in the throat, basally connected by a thin membrane
(false corona) which is entire to deeply incised between the filaments. Ovary with 1-9
362
Flora Malesiana ser. I, Vol. 11 (2) (1993)
ovules near the bottom of each cell; style filiform; stigma entire and small. Fruits fleshy,
finally rupturing laterally. Seeds spongy, green.
Distribution — About 50 species in tropical and subtropical America, a few cultivated;
in Malesia there is one naturalized species.
Hymenocallis littoralis (Jacq.) Salisb., Trans.
Hort Soc. Lond. 1 (1812) 338; Merr., Sp. Blanc.
(1918) 98; Enum. Philipp. How. PL 1 (1923)
211; Backer & Bakh. f., FL Java 3 (1968) 138;
Peekel, FL Bismarck Arch. Natur. (1984) 89.
— Pancratium littorale Jacq., Select. Stirp.
Amer. Hist. (1763) 99; Ker-GawL, Curt. Bot.
Mag. 21 (1805) t. 805; Safford, Contr. U.S.
Nat. Herb. 9 (1905) 342. — Type: Jacquin,
West Indies.
Hymenocallis adnata Herbert, Amaryll. Suppl.
(1837) 215, nom. illeg.; Naves, Nov. App.
(1880) 256.
Hymenocallis tenuiflora Herbert, Amaryll. Suppl.
(1837) 215; Naves, Nov. App. (1880) 149. —
Type: Hilton 143, New Granada.
Pancratium illyricum auct. non L.: Blanco, FL
Filip. ed. 3, 1 (1877) 316.
Pancratium maritimum auct. non L.: Blanco, FL
Filip. ed. 3 (1877) 316; Naves, Nov. App.
(1880) 256.
Bulb spherical, 7.5-10 cm in diameter. Leaves
2-ranked, stalkless, oblong, 40-120 cm by 1.5-5
(-7) cm, acute. Inflorescences 6-11-flowered; pe-
duncle 45-60 cm long; spathes 6-7 cm long; pedi-
cel none to very short Flowers fragrant. Perigone-
tube 14-17 cm long; segments linear, channelled,
7.5-12 cm long and 2-3 mm wide, tipped. False
corona 2.5-3 cm high; free part of filaments c. 6
cm. Ovules 4-5 per cell, style slighdy exceeding
the stamens.
Distribution - South to Central America; culti-
vated and naturalized in tropical Africa, tropical
Asia, Malesia and the Pacific Islands. Malesia: Java,
Philippines (Luzon), Bismarck Archipelago.
Habitat - Along the seashore and in moist sandy
places at low altitudes.
CULTIVATED SPECIES
Hymenocallis caribaea (L.) Herbert — Backer
& Bakh. f., FL Java 3 (1968) 138.
Native of the West Indies; cultivated as an orna-
mental. Flowers fragrant; perigone greenish; false
corona 2-3 cm long.
Hymenocallis x macrostephana Baker —
Backer & Bakh. f., FL Java 3 (1968) 138.
A hybrid between the two following species;
cultivated as an ornamental. Flowers fragrant; peri-
gone white to greenish; false corona 5.5-6 cm
long.
Hymenocallis narcissiflora (Jacq.) MacBr.
— Backer & Bakh. f., FL Java 3 (1968) 138.
Native of South America; cultivated as an orna-
mental. Rowers fragrant; perigone green and white;
false corona 6-7.5 cm long.
Hymenocallis speciosa (Salisb.) Salisb. —
Backer & Bakh. f., FL Java 3 (1968) 138.
Native of the West Indies; cultivated as an or-
namental. Rowers fragrant; perigone greenish and
white; false corona 2.5-5 cm long.
PANCRATIUM
Pancratium L., Sp. PI. (1753) 290. — Type species: Pancratium maritimum L.
Bulbous herbs. Leaves radical, biseriate, sessile. Inflorescences umbellate, few- to
1 -flowered, with 1 or 2 spathes. Flowers white. Tepals subequal, united at the base into a
long and narrow tube. Stamens inserted in the throat, basally connected by a thin mem-
brane (false corona) which is sometimes 1- or 2-toothed between the filaments; anthers
dorsifixed. Ovary with 10 or more biseriate ovules filling each cell; style filiform; stigma
entire. Capsules loculicidal, 3-valved. Seeds angular by pressure, black. — Fig. 2.
Distribution — About 15 species in S Europa, tropical Africa and Asia, a few culti-
vated; in Malesia one indigenous species and a few others cultivated.
Geerinck — Amaryllidaceae
363
Pancratium zeylanicum L., Sp. PI. (1753)
290; Hooker, Curt. Bot. Mag. 52 (1825) L 2528;
Miq., Fl. Ned. Indie 3 (1859) 582; Blanco, Fl.
Filip., ed. 3, 1 (1877) 317; Naves, Nov. App.
(1880) 256; Baker, Handb. Amaryll. (1888) 48;
Koord., Meded. Lands Plantent. 19 (1898) 310;
Fig. 2. Pancratium zeylanicum L. Flower without
ovary (Korthals s.n., L 897.324-159).
Merr., Fl. Manila (1912) 148; Sp. Blanc. (1918)
99; Enum. Philipp. Flow. PI. 1 (1925) 212;
Backer & Bakh. f., Fl. Java 3 (1968) 141. —
Type: Probably from Sri Lanka, but perhaps of
garden origin.
Glabrous herbs. Leaves linear to narrowly ellip-
tic, acute, 12-40 cm by 8-30 mm. Inflorescences
1 -flowered; peduncle 4-15 cm long; spathe 1, 2-
nerved, apically bilobed, 4-5 cm long; pedicel ab-
sent or very short. Flowers with 2.5-3.5 cm long
perigone-tube, lobes linear, 6.5-8 cm long and
4-7 mm wide, acuminate. False corona toothed,
3.5-4 cm high; free part of filaments 2.5-4 cm.
Style exceeding the stamens. - Fig. 2.
Distribution - Malesia: Borneo (Pandaran),
Java, Philippines (Luzon, Palawan, Biliran), Cele-
bes (Minahasa), Moluccas (Ambon, Gorontala).
Also cultivated in gardens.
Habitat - Grasslands, light forest arid planta-
tions, up to 600 m altitude.
CULTIVATED SPECIES
Pancratium longiflorum Roxb. — Miq., Fl.
Ned. Indie 3 (1859) 583.
Doubtful species.
Pancratium maritimum L. — Backer &
Bakh. f., Fl. Java 3 (1968) 141.
Native to the Mediterranean region; cultivated
as an ornamental. Free part of the filaments 6-10
mm long.
Pancratium verecundum AiL — Backer &
Bakh. f., Fl. Java 3 (1968) 141.
Native of SE Asia; cultivated as an ornamental.
Free part of the filaments 2-3 cm long.
PROIPHYS
Proiphys Herbert, App. (1821) 42; Mabberley, Taxon 29 (1980) 597. — Eurycles Salisb.
ex Schult. in Roem. & Schult., Syst. Veget. 7/2 (1830) 909, nom. illeg., superfl. —
Type species: Proiphys amboinensis (L.) Herbert.
Bulbous herbs. Leaves radical, petioled, subcircular to elliptic, with numerous paral-
lel nerves. Inflorescences umbellate, many-flowered; spathes 2-4. Flowers pedicellate,
white. Tepals united into a narrow tube; segments subequal, erect to patent, elliptic to ob-
ovate, apiculate. Stamens inserted in the throat, basally connected by a membrane (false
corona) which is deeply bilobed between the filaments and shorter than the perigone;
anthers medifixed. Ovary with 2 ovules in each cell; style filiform; stigma entire, small.
Fruits dry or slightly fleshy, rupturing irregularly. Seeds globose, green. — Figs. 3, 4.
Distribution — In Australia 3 species, two of them also indigenous in Malesia.
364
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 3. Proiphys amboinensis (L.) Herbert. Cultivated in the Hortus Botanicus Leiden, 1961 (Photograph
Rijksherbarium Leiden).
Geerinck — Amaryllidaceae
365
Fig. 4. Proiphys amboinensis (L.) Herbert, a. Flower; b. leaf. — P. alba (R. Brown) Mabberley. c. Leaf
(a, b: de Boer bb 2225; c: McKee 1713).
KEY TO THE SPECIES
la. Leaves elliptic to ovate, less than 15 cm wide, cuneate at base. Spathes 2-3 cm long.
Peri gone-tube 8-15 mm long; segments less than 25 mm long 1. P. alba
b. Leaves subcircular to broadly ovate, slightly cordate at base, 15 cm or more wide.
Spathes 3-10 cm long. Perigone-tube 22-35 mm long; segments 25 mm long or
more 2. P. amboinensis
366
Flora Malesiana ser. I, Vol. 11 (2) (1993)
1. Proiphys alba (R. Brown) Mabberley, Taxon
29 (1980) 601. — Eurycles alba (R. Brown)
F. Muell., Sec. Cens. Austral. PL (1889) 196.
— Type: R. Brown, Queensland.
Bulb 2-4 cm diam. Leaves: petiole 7-35 cm
long; limb elliptic to ovate, cuneate at base, 10-
35 cm long and 2-10 cm wide, margin flat. Umbel
10-30-flowered; spathes 3 or 4, 2-8 cm long;
pedicels 7-30 mm long. Flowers with 8-15 mm
long perigone-tube, lobes 6-24 mm long. False
corona 5-12 mm long; free part of filaments 2-3
mm. Fruits unknown. - Fig. 4c.
Distribution - Australia; Malesia: New Guinea
(Merauke). Also cultivated in gardens.
Habitat - Woodlands at low altitudes.
2. Proiphys amboinensis (L.) Herbert, Appen-
dix (1821) 42. — Pancratium amboinense L.,
Sp. PL (1753) 291; Blanco, Fl. Filip. ed. 3, 1
(1877) 317. — Eurycles amboinensis (L.) Lind-
ley in Loud. Encycl. PL (1829) 242; Miq., Fl.
Ned. Indie 3 (1859) 582; Naves, Nov. App.
(1880) 255; Vidal, Phan. Cuming. Philipp.
(1885) 152; Rev. PL Vase. Filip. (1886) 276;
Koord., Meded. Lands Plantent. 19 (1898) 310;
Merr., Fl. Manila (1912) 149; Sp. Blanc. (1918)
99; Hallier, Nova Guinea 8 (1913) 900; Merr.,
Enum. Philipp. Row. PL 1 (1925) 211; Hen-
derson, Mai. Wild FL, Monocot. (1954) 166;
Backer & Bakh. f., Fl. Java 3 (1968) 140; Peekel,
Fl. Bismarck Arch. Natur. (1984) 89. — Type:
Indonesia, illustration in Commelin, Hort. Med.
Amstel. PL 1 (1697) t. 39, as Narcissus am-
boinensis.
Eurycles sylvestris Salisb. ex Schult., Syst. Veget.
7 (1830) 909, nom. illeg.; Salisb., Trans. Hort.
Soc. Lond. ed. 1, 1 (1812) 337, nomen; Ridley,
Fl. Mal.Penins. 6 (1924) 303. — Type: Rumph.,
Herb. Amb. 6, t. 70, as Caepa sylvestris.
Eurycles javanica Roemer, Prospect. Fam. Nat.
Syn. Monogr. 4 (1847) 157, nom. illeg.; Miq.,
Fl. Ned. Indie 3 (1859) 582.
Bulb to 8 cm diam. Leaves: petiole 15-60 cm
long; limb subcircular to broadly ovate, hardly cor-
date at base, emarginate to shortly apiculate, 20-
30 cm long and 15-35 cm wide, margin undulate.
Umbel 5-25-flowered; peduncle 15-90 cm long;
spathes 3 or 4, 3-10 cm long; pedicels to 4.5 cm
long. Flowers with 22-35 mm long perigone-
tube, lobes 25-40 mm long. False corona 7-10
mm long; free part of filaments 10-12 mm long.
Fruits 15-30 mm diam. - Figs. 3, 4a, b.
Distribution - Continental Asia (Thailand),
Australia; in Malesia: Philippines (Luzon, Mindo-
ro, Palawan), Celebes (Minahasa, Butung), Lesser
Sunda Islands (Bali, Timor), Moluccas (Ambon),
New Guinea (Merauke, Milne Bay, Fly River),
Bismarck Archipelago. Also cultivated in gardens.
Habitat - Seashores and rocky places, up to
500 m altitude.
Subfamily Hypoxidoideae
Inflorescence a raceme, occasionally head-like or flowers solitary, with bracts but never
spathes. Flowers actinomorphic. Tepals free or united into a tube. Stamens free or in-
serted on the perigone-tube.
Six genera.
CURCULIGO
Curculigo Gaertn., Fruct. Sem. PL 1 (1788) 63. — Type species: Curculigo orchioides
Gaertn.
Gethyllis L., Sp. PI. (1753) 442. — Type species: Gethyllis afra L.
Molineria Colla, Hort. Ripul., App. 2 (1826) 331. — Type species: Molineria plicata
Colla, nom. illeg.
Thickly rhizomatous herbs, hairy or glabrous. Leaves radical, often petiolate, linear to
broadly elliptic. Inflorescences racemes or spikes, sometimes head-like, many- to few-
flowered. Flowers with the tepals united in a very short to elongate tube. Stamens free,
Geerinck — Amaryllidaceae
367
inserted at the mouth of the perigone-tube. Fruits fleshy, indehiscent or irregularly dehis-
cent, often with a persistent beak. — Fig. 5.
Distribution — About 20 species in the tropics; in Malesia 5 indigenous species and
1 variety; perhaps one species cultivated.
KEY TO THE SPECIES AND VARIETIES
la. Leaves up to 20 cm long. Inflorescences lax, up to 8-flowered . 4. C. orchioides
b. Leaves usually longer than 20 cm. Inflorescences compact 2
2a. Inflorescences sessile or with scape to 4(-10) cm long. Fruit beaked 3
b. Inflorescences with scape 5-30 cm long. Fruit beaked or not 4
3a. Perigone-tube 8-15 mm long 3a. C. latifolia var. latifolia
b. Perigone-tube 20-40 mm long 3b. C. latifolia var. megacarpa
4a. Bracts glabrous or ciliate at the top. Perigone-tube 7-8 mm long. Fruit beaked
2. C. erecta
b . Bracts villous to glabrescent. Perigone-tube to 2 mm long. Fruit not beaked .... 5
5a. Inflorescences head-like and deflexed 1. C. capitulata
b. Inflorescences elongate and erect 5. C. racemosa
1. Curculigo capitulata (Lour.) Kuntze, Rev.
Gen. PI. 1 (1891) 703; Backer, Handb. Fl. Java
1 (1925) 104; Merr., Enum Philipp. Flow. PI.
1 (1925) 213. — Leucojum capitulatum Lour.,
Fl. Cochinch. (1790) 199. — Molineria capitu-
lata (Lour.) Herbert, Amaryll. (1837) 84; Backer
& Bakh.f., Fl. Java 3 (1968) 209. — Type:
unknown.
Curculigo recurvata Dryand. in Aiton, Hort. Kew.,
ed. 2, 2 (1814) 253; Miq., Fl. Ned. Indie 3
(1859) 585; Naves, Nov. App. (1880) 253;
Koord., Meded. Lands Plantent. 19 (1898) 310;
Ridley, Mat. Fl. Mai. Penins. 2 (1907) 65;
Koord., Exk. Fl. Java 1 (1911) 304; Elmer,
Leafl. Philipp. Boi. 5 (1913) 1645; Hallier,
Nova Guinea 8 (1913) 901; Laut., Bot. Jahrb.
50 (1913) 304; Ridley, Fl. Mai. Penins. 4
(1925) 299; Peekel, Fl. Bismarck Arch. Natur.
(1984) 91. — Molineria recurvata (Dryand.)
Herbert, Amaryll. (1837) 84. — Type: Hort.
Kew, Bangladesh.
Curculigo glabra Merr., Philipp. J. Sc, BoL 2
(1907) 267; Enum. Philipp. Row. PI. 1 (1925)
213. — Type: Merrill 5750, Mindoro.
Hairy herb. Leaves long-petiolate, elliptic to
broadly elliptic, 60-150 cm long and 5-15 cm
wide, nearly glabrous. Inflorescence a deflexed and
head-like raceme, 2.5-7 cm long and wide; pedun-
cle 7-30 cm long, tomentose to glabrescent at
fructitication: bracts brownish, 1.5-5 cm long.
villous to glabrescent at fructification. Flowers
subsessile, yellow, villous. Perigone-tube 1-2
mm long; lobes 6-8 mm long. Fruits whitish to
green, 10-15 mm long, not beaked. - Fig. 5a.
Distribution - Continental Asia (Nepal, India,
Sri Lanka, Bangladesh, Thailand), Taiwan, Aus-
tralia, Pacific Islands (Solomon Islands, Hawaii);
Malesia: Sumatra, Malaya, Singapore, Java, Phi-
lippines (Luzon, Negros, Biliran, Mindanao),
Celebes, Moluccas (Halmaheira, Talaud, Ternate,
Ambon), New Guinea, Manus I.
Habitat - Primary and secondary forests, from
sea-level up to c. 2000 m altitude.
Uses - The fruits are edible. Leaf fibres are (or
were) used in the Philippines, according to Burkill,
Diet. Econ. Prod. Malay Penins. 1 (1935) 703.
2. Curculigo erecta LauL, Bot. Jahrb. 50 (Aug.
1913) 304. — Type: Schulze 179, Papua New
Guinea, Sepik River.
Curculigo scapigera Hallier, Nova Guinea 8 (Sept.
1913) 901. — Type: Versteeg 1084, New Guinea.
Hairy herb. Leaves to 120 cm long and 6 cm
wide. Inflorescences head-like, compactly ovoid to
cylindrical, 2-4 cm long and 1-2 cm wide; pedun-
cle tomentose to glabrescent, 5-30 cm long; bracts
glabrous or ciliate at the top, to 2 cm long and 8
mm wide; pedicels c. 6 mm long. Flowers yellow.
Perigone-tube 7-8 mm long. Fruits 10-18 mm
long, beaked. - Fig. 5 b.
368
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Geerinck — Amaryllidaceae
369
Distribution - Malesia: E Sumatra, Philippines
(Luzon), New Guinea (Irian Jaya). Not (yet) seen
from Borneo and Celebes.
Habitat - Primary forest, montane forest, also
in swamps. Altitudinal range uncertain.
3. Curculigo latifolia Dryand. in Aiton, Hort.
Kew. ed. 2, 2 (1811) 253; Miq., Fl. Ned. Indie
3 (1859) 595; Ridley, Mat. Fl. Mai. Penins.
2 (1907) 66; Fl. Mai. Penins. 4 (1925) 300;
Backer, Handb. Fl. Java 1 (1925) 104; Hender-
son, Mai. Wild Fl., Monocot. (1954) 167. —
Molineria latifolia (Dryand.) Herbert ex Kurz,
Nat. Tijd.. Ned. Indie 27 (1864) 232; Backer &
Bakh. f., Fl. Java 3 (1968) 209. — Type: Hort.
Kew, India.
Curculigo glabrescens (Ridley) Merr., J. Str. Br.
Roy. As. Soc. 85 (1922) 163. — Curculigo
latifolia var. glabrescens Ridley, Mat. Fl. Mai.
Penins. 2 (1907) 67. — Type: Ridley, Malaya.
Curculigo sumatrana Roxb., Fl. Ind. 2 (1824) 146;
Ridley, Mat. Fl. Mai. Penins. 2 (1907) 66; Fl.
Mai. Penins. 4 (1925) 300 — Molineria suma-
trana (Roxb.) Herbert, Amaryll. (1837) 84. —
Type: Hort. Kew, India.
Molineria plicata Colla, Hon. Ripul., App. 2 (1826)
331, pi. 18, nom. illeg.; Kurz in Miq., Ann.
Mus. Bot. Lugd.-Bat. 4 (1869) 176. — Type:
Sumatra.
Curculigo villosa Wall, ex Kurz in Miq., Ann. Mus.
Bot. Lugd.-Bat. 4 (1869) 176; Ridley, Fl. Mai.
Penins. 4 (1925) 300. — Curculigo latifolia
var. villosa (Wall, ex Kurz) Baker, J. Linn. Soc.
17 (1878) 125. — Type: Wallich 5163A, Sin-
gapore.
Curculigo agusanensis Elmer, Leafl. Philipp. Bot.
5 (1913) 1645; Merr., Enum. Philipp. Flow.
PI. 1 (1925) 213. — Types: Elmer 13584, Min-
danao, Mt Urdaneta.
Curculigo weberi Elmer, Leafl. Philipp. Bot. 5
(1913) 1646; Merr., Enum. Philipp. Row. PI.
1 (1925) 213. — Type: Elmer 14304, Minda-
nao, Mt Urdaneta
Curculigo brevipedunculata Elmer, Leafl. Philipp.
BoL 5 (1918) 1647; Merr., Enum. Philipp.
Flow. PI. 1 (1925) 213. — Type: Elmer 13164,
Palawan, Mt Pulgar.
Curculigo borneensis Merr., J. Str. Br. Roy. As.
Soc. 85 (1922) 162. — Type: Ramos 1712,
Borneo, Sandakan.
Hairy herb. Leaves long-petiolate, elliptic to
broadly elliptic, 30-100 cm long and 5-10 cm
wide, glabrous or nearly so. Inflorescences ovoid
to cylindrical, compact, 2-6 cm long and wide,
sessile or peduncle to 4(-10) cm long, glabrescent;
bracts green, glabrous or ciliate at margins, 1-6
cm long. Flowers subsessile, yellow, villous.
Perigone-tube 8-40 mm long; lobes 8-12 mm
long. Fruits 10-25 mm long, beaked, white to
green.
a. var. latifolia
Perigone-tube 8-15 mm long. - Fig. 5c.
Distribution - Continental Asia (India, Burma,
Thailand); Malesia: Sumatra, Lingga, Bangka, Ma-
laya (Pahang), Borneo (Sarawak, Sabah, Kaliman-
tan), Java, Philippines (Palawan, Balabac, Samar),
Celebes.
Habitat - Forests, at high altitudes.
Uses - The fruits are edible. Burkill, Diet. Econ.
Prod. Mai. Penins. 1 (1935) 704 and Heyne, NutL
PI. Indonesia 1 (1950) 451 report the (former) use
of rolled leaves as string and of extracted leaf-fibres
for fishing nets etc. in Borneo and Malaya. Medic-
inal uses are also mentioned for Malaya (Burkill,
I.e.).
Note - The specimen Jacobs 5549 from Borneo
shows three subsessile inflorescences and one with
a peduncle 10 cm long.
b. var. megacarpa (Ridley) Geerinck, comb,
nov. — Curculigo megacarpa Ridley, J. Str. Br.
Roy. As. Soc. 41 (1904) 33; Mat. Fl. Mai.
Penins. 1 (1907) 67; Fl. Mai. Penins. 4 (1925)
300. — Type: Ridley, Malaya, Perak, Thaiping
Hills.
Perigone-tube 20-40 mm long.
Distribution - Continental Asia (Thailand);
Malesia: Malaya (Perak), Borneo (Sandakan), Java
(also Panaitan I.).
Habitat - Rain forest, at high altitudes.
Note - Further material is necessary to confirm
the rank of this taxon.
Fig. 5. Curculigo capitulata (Lour.) Kuntze. a. Inflorescence. — C. erecta Laut. b. Inflorescence. — C.
latifolia Dryand. var. latifolia. c. Inflorescence. — C. orchioides Gaertn. d. Flowering plant. — C. race-
mosa Ridley, e. Inflorescence (a: de Wilde & de Wilde-Duyfjes 13539; b: Brass 6994; c: Geesink &
Hattink6401; ±7ippel272; e: Jacobs 5610).
370
Flora Malesiana ser. I, Vol. 11 (2) (1993)
4. Curculigo orchioides Gaertn., Fruct. Sem.
PI. 1 (1788) 63; Miq., Fl. Ned. Indie 3 (1859)
585; Backer, Handb. Fl. Java 1 (1925) 103;
Merr., Enum. Philipp. Flow. PI. 1 (1925) 213;
Backer & Bakh. f., Fl. Java 3 (1968) 209. —
Type: unknown.
Gethyllis acaulis Blanco, Fl. Filip. (1837) 260;
Merr., Dep. Int. Bur. Gov. Labor. 27 (1905) 85.
— Type: Blanco, Philippines, Manila.
Hairy herb. Leaves subpetiolate, linear to ellip-
tic, to 20 cm long, 8 cm wide. Inflorescence lax,
up to 8-flowered; bracts long-acuminate, 2-3.5 cm
long, long-ciliate; pedicels 2-3 cm long. Flowers
bright yellow. Perigone-tube 5-20 mm long; lobes
c. 6.5 mm long. Fruit green, beaked. - Fig. 5d.
Distribution - Continental Asia (India, Laos,
Cambodia, Vietnam, Thailand), Taiwan, Pacific
Islands (Marianes, New Caledonia); Malesia: Su-
matra, Borneo (Kalimantan), Java, Philippines
(Luzon, Samar, Mindanao), Celebes, Lesser Sunda
Islands (Sumba, Flores, Timor), Moluccas (Am-
bon), New Guinea, Misima I.
Habitat - Grassland, at low altitudes.
Uses - Medicinal use of roots is mentioned by
Burkill (Diet. Econ. Prod. Malay Penins. 1, 1935,
703) and Quisumbing (Medic. PI. Philipp. 1951,
172) on the strength of Indian literature.
5. Curculigo racemosa Ridley, J. Str. Br. Roy.
As. Soc. 44 (1805) 198. — Type: Ridley, Bor-
neo, Mt Matang.
Hairy herb. Leaves 50-100 cm long and 8-10
cm wide. Inflorescence elongate and erect; peduncle
(10-)20-40 cm, tomentose; bracts to 2.5 cm long,
3 mm wide, villous; pedicels 8-15 mm long.
Flowers bright yellow. Perigone-tube 1-2 mm
long; lobes 6-12 mm long and 1.5-2 mm wide.
Fruits green, 8-12 mm long, not beaked. - Fig. 5e.
Distribution - Malesia: Borneo (Sarawak, Sabah,
Kalimantan).
Habitat - Rain forest, old secondary forest, at
high altitudes.
Note - The specimen Jacobs 5610 is said to be
an epiphyte on a hollow tree trunk.
CULTIVATED SPECIES
Curculigo ensifolia R. Br. — LauL, Bol Jahrb.
50(1913)304.
Native of Australia: perhaps a synonym of C.
orchioides Gaertn.
HYPOXIS
Hypoxis L., Syst. Nat., ed. 10 (1789) 986.
Hairy or glabrous herbs, bulbous or thickly rhizomatous. Leaves radical, linear to nar-
rowly elliptical. Inflorescence a 1- to many-flowered raceme; bracts narrow. Flowers with
free tepals. Stamens free. Fruit a dry capsule, regularly dehiscent. — Fig. 6.
Distribution — About 100 species, predominantly in tropical Africa; one indigenous
species in Malesia and one cultivated.
Hypoxis aurea Lour., Fl. Cochinch. (1790)
200; Ridley, Mat. Fl. Mai. Penins. 1 (1907)
64; Koord., Exk. Fl. Java 1 (1911) 305; LauL,
BoL Jahrb. 50 (1913) 305; Backer, Handb. Fl.
Java 1 (1925) 105; Merr., Enum. Philipp. Row.
PL 1 (1925) 214; Backer & Bakh. f., Fl. Java 3
(1968) 209. — Type: Loureiro, Cochinchina,
So Koung.
Hypoxis franquevillei Miq., Fl. Ned. Indie 3 (1859)
586; Naves, Nov. App. (1880) 253. — Type:
Franqueville, Java, Penangoen.
Plant hairy; rootstock globose to elongate.
Leaves linear to narrowly elliptic, acuminate, 8-25
cm long and 1-4 mm wide. Inflorescences 4-8
cm long, shorter than the leaves, 1- to few-flower-
ed; bracts linear; pedicels 1.5-4 mm long. Flowers:
tepals 5-6 mm long and about 1.5 mm wide. Cap-
sules about 7 mm long. - Fig. 6.
Distribution - Continental Asia (China, India,
Cambodia, Thailand), Japan, Taiwan; Malesia: Su-
matra, Malaya, Java, Philippines (Luzon, Minda-
nao), Celebes, Flores, New Guinea (Irian Jaya).
Habitat - Grassland and other sunny places, not
at very low altitudes.
CULTIVATED SPECIES
Hypoxis hygrometrica Labill. — Laut., BoL
Jahrb. 50 (1913) 305.
Native of Australia; cultivated in New Guinea.
Plant sparsely hairy; inflorescence 1-2-flowered.
Geerinck — Amaryllidaceae
371
Fig. 6. Hypoxis aurea Lour. Flowering plant (Brass
11783).
CULTIVATED SPECIES
OF NON-MALESIAN GENERA
Amaryllis belladonna L. — Backer, Handb.
Fl. Java 1 (1925) 85; Steiner, Philipp. Ornam.
PI. (1960) 136; Backer & Bakh. f., Fl. Java 3
(1968) 133.
Native of South Africa; cultivated as an orna-
mental. Umbel 6-12-flowered; perigone dark red
or pink to white.
Clivia miniata (Lindley) Bosse — Backer,
Handb. Fl. Java 1 (1925) 84; Backer & Bakh. f.,
Fl. Java 3 (1968) 133.
Native of South Africa; cultivated as an orna-
mental. Umbel 12-20-flowered; perigone usually
orange.
Cooperia drummondii Herbert — Backer,
Handb. Fl. Java 1 (1925) 87; Backer & Bakh. f.,
Fl. Java 3 (1968) 135.
Native of N America; cultivated as an ornamen-
tal. Perigone white, often tinged with violet.
Cyrtanthus mackenii Hook. f. — Backer,
Handb. Fl. Java 1 (1925) 90; Backer & Bakh. f.,
Fl. Java, 3 (1968) 137.
Native of South Africa; cultivated as an orna-
mental. Umbel 4-10-flowered; flowers fragrant;
perigone 3.5-5 cm long, white to yellowish.
Cyrtanthus pallidus Sims — Backer, Handb.
Fl. Java 1 (1925) 90; Backer & Bakh. f., Fl.
Java 3 (1968) 137.
Native of South Africa; cultivated as an orna-
mental. Umbel 4-7-flowered; perigone pale pink,
4-5.5 cm long.
Eucharis amazonica Linden ex Planch. —
Backer, Handb. Fl. Java 1 (1925) 92; Backer &
Bakh. f., Fl. Java 3 (1968) 139. — Eucharis
grandiflora auct non Planch. & Linden: Merr.,
Enum. Philipp. Row. PI. 1 (1925) 211; Peekel,
Fl. Bismarck Arch. Natur. (1984) 89.
Native of Colombia; cultivated as an ornamen-
tal. Leaves several per bulb; umbel 4-8-flowered;
flowers fragrant; false corona white with greenish
or yellowish stripes.
Eucharis Candida Planch. & Linden — Koord.,
Exk. Fl. Java 1 (1911) 303; Backer, Handb. Fl.
Java 1 (1925) 92; Backer & Bakh. f., Fl. Java 3
(1968) 139.
Native of Colombia; cultivated as an ornamen-
tal. Leaf 1 per bulb; umbel 6- 10- flowered; flowers
fragrant; false corona yellow.
372
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Eucharis mastersii Baker — Eucharis gran-
diflora auct. non Planch. & Linden: Koord.,
Exk. Fl. Java 1 (1911) 303; Backer, Handb. Fl.
Java 1 (1925) 92; Steiner, Philipp. Ornam. PI.
(1960) 137; Backer & Bakh. f., Fl. Java 3 (1968)
139.
Native of Colombia; cultivated as an ornamen-
tal. Leaves several per bulb; umbel 2-5-flowered;
flowers fragrant in the evening; false corona green
and yellow.
Eucharis sanderi Baker — Backer, Handb. Fl.
Java 1 (1925) 92; Backer & Bakh. f., Fl. Java 3
(1968) 139.
Native of Colombia; probably a hybrid between
the two preceding species; perhaps cultivated as an
ornamental. Umbel 2-3-flowered.
Eucharis subedentata Baker — Backer, Handb.
Fl. Java 1 (1925) 92; Backer & Bakh. f., Fl.
Java 3 (1968) 139.
Native of Colombia; cultivated as an ornamen-
tal. Leaves several per bulb; umbel 6-8-flowered;
false corona not toothed.
Griffmia ornata Moore — Backer, Handb. Fl.
Java 1 (1925) 84; Backer & Bakh. f., Fl. Java 3
(1968) 133.
Native of Brazil; cultivated as an ornamental.
Umbels 10-16-flowered; perigone violet.
Hippeastrum advenum Ker-Gawl. — Backer,
Handb. Fl. Java 1 (1925) 96; Backer & Bakh. f.,
Fl. Java 3 (1968) 142.
Native of Chile; cultivated as an ornamental.
Perigone-lobes 3.5-4.5 cm long, red, pink or yel-
low.
Hippeastrum aulicum (Ker-Gawl.) Herbert —
LauL, Bol Jahrb. 50 (1913) 303; Backer, Handb.
Fl. Java 1 (1925) 96; Backer & Bakh. f., Fl.
Java 3 (1968) 142; Peekel, Fl. Bismarck Arch.
Natur. (1984) 89.
Native of Brazil; cultivated as an ornamen-
tal. Perigone-lobes 9-15 cm long, red with darker
veins.
Hippeastrum pra tense (Poepp.) Baker —
Backer, Handb. Fl. Java 1 (1925) 96; Backer &
Bakh. f., Fl. Java 3 (1968) 142.
Native of Chile; cultivated as an ornamental.
Perigone-lobes 6-6.5 cm long, bright red to pur-
plish.
Hippeastrum puniceum (Lam.) Kuntze —
Backer & Bakh. f., Fl. Java 3 (1968) 143. —
Hippeastrum equestre (Ait.) Herbert: Koord.,
Exk. Fl. Java 1 (1911) 303; Backer, Handb. Fl.
Java 1 (1925) 97; Steiner, Philipp. Ornam. PI.
(1960) 137.
Native of Central America and West Indies; cul-
tivated as an ornamental. Perigone-lobes red to
orange-red and greenish to whitish at base.
Hippeastrum reginae (L.) Herbert — Backer,
Handb. Fl. Java 1 (1925) 97; Backer & Bakh. f.,
Fl. Java 3 (1968) 142. — Amaryllis atamasco
auct. non L.: Blanco, Fl. Filip. (1837) 254. —
Hippeastrum miniatum auct. non Herbert Merr.,
Enum. Philipp. Flow. PI. 1 (1925) 212.
Native of tropical America; cultivated as an or-
namental. Perigone-lobes 8-12 cm long, red with
greenish 5 -pointed star at base.
Hippeastrum reticulatum (L'Her.) Herbert —
Laut., Bol Jahrb. 50 (1913) 302; Backer, Handb.
Fl. Java 1 (1925) 97; Steiner, Philipp. Ornam.
PI. (1960) 137; Backer & Bakh. f., Fl. Java 3
(1968) 142; Chin, Malays. Fl. Col. (1974) 145;
Peekel, Fl. Bismarck Arch. Natur. (1984) 87.
Native of Brazil; cultivated as an ornamental.
Perigone-lobes 10-12.5 cm long, red-mauve with
darker veins.
Hippeastrum rutilum Ker-Gawl. — Hippeas-
trum splendens Herbert: Backer, Handb. Fl. Java
1 (1925) 97; Backer & Bakh. f., Fl. Java 3
(1968) 142.
Native of Brazil; cultivated as an ornamental.
Perigone-lobes 8-12 cm long, greenish and api-
cally red.
Hippeastrum stylosum Herbert — Backer,
Handb. Fl. Java 1 (1925) 97; Backer & Bakh. f.,
Fl. Java 3 (1968) 142.
Native of Guyana and Brazil; cultivated as an
ornamental. Perigone-lobes c. 10 cm long, bright
red with a pale median band.
Hippeastrum vittatum (L'Her.) Herbert —
Backer, Handb. Fl. Java 1 (1925) 97; Backer &
Bakh. f., Fl. Java 3 (1968) 299.
Native of Peru and Brazil; cultivated as an or-
namental. Perigone-lobes 7.5-15 cm long, white
with red stripes.
Leucojum aestivum L. — Backer & Bakh. f.,
Fl. Java 3 (1968) 144.
Native to S Europe; cultivated as an ornamen-
tal. Umbel 3-5(-7)-flowered; perigone-lobes 1-1.5
cm long.
Geerinck — Amaryllidaceae
373
Narcissus jonquilla L. — Backer, Handb. Fl.
Java 1 (1925) 94; Backer & Bakh. f., Fl. Java
3 (1968) 140. — Narcissus pseudonarcissus
auct. non L. (?): Koord, Exk. Fl. Java 1 (1911)
303.
Native of S Europe; cultivated as an ornamen-
tal. Umbel 2-5-flowered; flowers fragrant; peri-
gone green and yellow.
Narcissus tazetta L. — Backer, Handb. Fl. Java
1 (1925) 94; Backer & Bakh. f., Fl. Java 3
(1968) 140. — Narcissus poeticus auct. non L.
(?): Koord., Exk. Fl. Java 1 (1911)303.
Native to S Europe and temperate Asia; cul-
tivated as an ornamental. Umbel 3-20-flowered;
perigone whitish.
Scadoxus multiflorus (Martyn) Raf. — Hae-
manthus multiflorus Martyn: Backer, Handb.
Fl. Java 1 (1925) 83; Steiner, Philipp. Ornam.
PI. (1960) 137; Backer & Bakh. f., Fl. Java 3
(1968) 132; Chin, Malays. Fl. Col. (1974) 143.
— Haemanthus puniceus auct. non L.: Koord.,
Exk. Fl. Java 1 (1911)302.
Native of tropical Africa; cultivated as an orna-
mental. Umbel spherical or hemispherical; peri-
gone red to pink.
Sprekelia formosissima (L.) Herbert —
Backer, Handb. Fl. Java 1 (1925) 96; Backer &
Bakh. f., Fl. Java 3 (1968) 141.
Native of Mexico and Guatemala; cultivated as
an ornamental. Perigone usually dark red, 9-12 cm
long. Poisonous.
Vallota speciosa (L. f.) Dur. & Schinz —
Backer, Handb. Fl. Java 1 (1925) 85; Backer &
Bakh. f., Fl. Java 3 (1968) 133.
Native of South Africa; cultivated as an orna-
mental. Umbel 4-9-flowered; perigone red to
white.
Zephyranthes atamasco (L.) Herbert — Backer,
Handb. Fl. Java 1 (1925) 86; Steiner, Philipp.
Ornam. PI. (1960) 138; Backer & Bakh. f., Fl.
Java 3 (1968) 135.
Native of N America; cultivated as an orna-
mental. Perigone about 7.5 cm long, white.
Zephyranthes brachyandra (Baker) Backer —
Backer & Bakh. f., FL Java 3 (1968) 134.
Native of Paraguay. Cultivated as an ornamen-
tal. Perigone 3-3.5 cm long, violet.
Zephyranthes Candida (Lindley) Herbert —
Backer, Handb. Fl. Java 1 (1925) 86; Backer &
Bakh. f., Fl. Java 3 (1968) 135; Peekel, Fl.
Bismarck Arch. Natur. (1984) 87.
Native of S America; cultivated as an orna-
mental and subnaturalized. Perigone about 1.5 cm
long, white slightly tinged with red.
Zephyranthes carinata Herbert — Backer,
Handb. Fl. Java 1 (1925) 86; Steiner, Philipp.
Ornam. PI. (1960) 138; Backer & Bakh. f., Fl.
Java 3 (1968) 134.
Native of S America; cultivated as an ornamen-
tal. Perigone 2.5-4.5 cm long, light pink.
Zephyranthes citrina Baker — Backer, Handb.
Fl. Java 1 (1925) 85; Backer & Bakh. f., Fl.
Java 3 (1968) 134.
Native of British Guyana; cultivated as an orna-
mental. Perigone 2-2.5 cm long, bright yellow.
Zephyranthes grandiflora Lindley — Backer,
Handb. Fl. Java 1 (1925) 86; Backer & Bakh. f.,
Fl. Java 3 (1968) 135. — Zephyranthes carinata
Herbert: Backer, Handb. Fl. Java 1 (1925) 86;
Backer & Bakh. f., Fl. Java 3 (1968) 134.
Native of Central America; cultivated as an or-
namental. Perigone 6-7.5 cm long, light pink.
Zephyranthes mesochloa auct. non Herbert (?):
Koord., Meded. Lands Plantent. 19 (1898) 311.
Zephyranthes rosea (Sprengel) Lindley —
Backer, Handb. Fl. Java 1 (1925) 86; Koord.,
Exk. Fl. Java 1 (1911) 302; Laut., Bot. Jahrb.
50 (1913) 302; Steiner, Philipp. Ornam. PI.
(1960) 138; Backer & Bakh. f., Fl. Java 3 (1968)
134; Peekel, Fl. Bismarck Arch. Natur. (1984)
85. — Atamasco rosea (Sprengel) Greene:
Merr., Enum. Philipp. Flow. PL 1 (1925) 210.
Native of Cuba. Cultivated as an ornamental and
subnaturalized. Perigone 3-3.5 cm long, pink.
Zephyranthes tubispatha (L'Her.) Herbert —
Backer, Handb. Fl. Java 1 (1925) 87; Backer &
Bakh. f., Fl. Java 3 (1968) 135; Peekel, Fl.
Bismarck Arch. Natur. (1984) 87.
Native of Central and South America; cultivated
as an ornamental. Perigone 4-5 cm long, white.
tinged with green.
Zephyranthes verecunda Herbert — Backer,
Handb. Fl. Java 1 (1925) 86; Backer & Bakh. f.,
Fl. Java 3 (1968) 135.
Native of Mexico; cultivated as an ornamental.
Perigone 3.5-5 cm long, white, tinged with pink.
Flora Malesiana ser. I, Vol. 1 1 (2) (1993) 375-384
ALLIACEAE
(J.R.M. Buijsen 1 , Leiden, The Netherlands)
Alliaceae Agardh, Theor. Syst. PI. (1858) 32; Dahlgren, Clifford & Yeo, Fam. Monocot.
(1985) 193-196. — Liliaceae subfam. Allioideae, Melchior in Melchior, Syll. Pflan-
zenfam. ed. 12(1964)521.
Perennial herbs with bulbs, bulb-like conns or rhizomes. Leaves simple, basally concen-
trated, spirally set or distichous. Inflorescence usually umbellate and with 1, 2 or more
membranous spathes. Flowers generally bisexual, actinomorphic or sometimes zygomor-
phic. Tepals in 2 whorls, free or often connate, forming a campanulate or tubular perianth
with erect, spreading or sometimes recurved lobes. Stamens usually 6 or sometimes 5
with several staminodes, free, inserted at the base of the tepals or in the perigone-tube;
anthers dorsifixed. Ovary superior, 3-celled, with axillary placentas, septal nectary grooves
present on the ovary; ovules 2 to several per locule. Fruit a loculicidal capsule. Seeds
often half-ovoid, half-globose or tetrahedral and triangular in transection, sometimes ovoid
or ellipsoid to subglobose and rounded in transection.
Distribution — As circumscribed by Dahlgren et al. (1985) this segregate from Lilia-
ceae s.l. comprises the South African Agapanthoideae, the mainly Chilean Gilliesioideae,
and Allioideae with the neogeic tribe Brodiaeeae and the nearly cosmopolitan Allieae.
Taxonomy — The taxonomic position of the genus Allium and related genera is still
disputed. Earlier botanists (e.g., Engler, Bentham, and Hooker) placed them in the Lilia-
ceae, as recently followed again by, e.g., Cronquist (1981) and Mabberley (1987). Alter-
natively, they were included in the Amaryllidaceae by, e.g., Hutchinson and Traub (see
Hanelt 1990). Often also Allium and its close relatives are recognized as a distinct family
Alliaceae, close to the Amaryllidaceae (Dahlgren et al. 1985). This controversy does not
matter much for Malesia since only Allium and Nothoscordum are found there, both not
indigenous.
References: Cronquist, A., Integrated system of classification of Flowering Plants (1981) 1208. —
Dahlgren, R.M.T., H.T. Clifford & P.F. Yeo, The families of the Monocotyledons (1985) 193. — Hanelt,
P., in H.D. Rabinowiich & J.L. Brewster (eds.), Onions and allied crops, Vol. 1 (1990) 2. — Mabberley,
D.J., The Plant-Book (1987) 331.
Palynology — The approximately 34 genera that make up the Alliaceae (see Dahlgren
& Clifford 1982) are palynologically rather poorly known. Pollen of representatives of 20
genera has been described (Tissot 1990), but nearly always with light microscopy only.
The most comprehensive works include Heusser (1971) and Schulze (1980). Beug (1961),
!) The International Board for Plant Genetic Resources IBPGR financed a taxonomic survey of Allium
species cultivated in SE Asia, that included a collection trip to Java in 1989.
(375)
376 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Radulescu (1973), Kuprianova & Aliev (1979) and Pastor (1981) give more or less ex-
tensive accounts on Allium pollen. Pollen of Nothoscordum, the other genus found in
Malesia, is dealt with by Schulze (I.e.).
Pollen grain size (length of largest equatorial axis) in Alliaceae is usually 25-55 um.
Several genera have significantly larger pollen: Agapanthus (up to 75 urn), Brodiaea (up to
78 urn), Dichelostemma (up to 70 urn), Milla (69-1 15 urn) and Triteleia (up to 72 urn).
The aperture system is always monosulcate. The sulcus is nearly as long as the long equa-
torial axis and restricted to the distal grain side, or it may continue on both ends on the
proximal side. Exine thickness is usually 1-2 urn, rarely < 1 urn or 2-3 urn. The exine is
columellate, and tectate or semitectate. The sexine is always thicker than the nexine. Orna-
mentation varies from psilate with (very) small perforations to reticulate. The diameter of
the perforations/lumina is mostly 0.5-1.5 um. Reticulate ornamentation with larger lumina
is found in pollen of Dichelostemma (1.5-4 um), Milla (3-7 um) and Muilla (1.5-4 um).
Reticulate ornamentation is usually heterobrochate, i.e. with lumina of different sizes
mingled on the proximal grain side. Towards the sulcus lumen size more or less gradually
decreases. Finely striate-rugulate ornamentation with minute perforations was found in
Allium using scanning electron microscopy (Pastor 1981).
The Alliaceae is a stenopalynous family. Palynologically the subfamilies are not dis-
tinct from each other. In the Allioideae two groups might be distinguished (Schulze I.e.).
On infrageneric level the length of the sulcus may sometimes be of taxonomic significance,
for example in Allium (Kuprianova & Aliev I.e.; Pastor I.e.). Pollen like that of the Allia-
ceae occurs in many other monocot families.
References: Beug, H.-J., Leitfaden der Pollenbestimmung (1961). — Dahlgren, R.M.T. & H.T. Clif-
ford, The monocotyledons: a comparative study (1982). — Heusser, C. J., Pollen and spores of Chile (1971).
— Kuprianova, L.A. & T. A. Aliev, BoL Zhurn. 64 (1979) 1273-1284. — Pastor, J., BoL Macaronesica
8/9 (1981) 189-214. — Radulescu, D., Acta Bot. Hon. Bucur. 1972-73 (1973) 133-248. — Schulze, W.,
Wiss. Z. Friedr.-Schiller-Univ. Jena, Math.-Naturwiss. R. 29 (1980) 595-606. — Tissot, C, Sixth bib-
liographic index to the pollen morphology of Angiosperms (1990).
R.W.J.M. van der Ham
Phytochemistry (compare also the relevant chapter under Amaryllidaceae) — All
members of Allium emit after wounding characteristic odours known as 'onion odour' and
'garlic odour'. Everywhere mankind met species of Allium, it made use of their spicy,
culinary and medicinal properties. Allium taxa, including a lot of cultivars of the onion
group (A. cepa), the garlic group (A. sativum) and the leek group (A. porrum), are
cultivated from time immemorial in southern Europe and the Near East. Allium kurrat
seems to have been taken in cultivation in ancient Arabia, Palestina and Egypt and A.
chinense, fistulosum, macrostemon and tuberosum had or still have many ancient
cultivars from India to China and Japan. Chives (A. schoenoprasum) were taken in
cultivation in postroman time in Europe.
There are three classes of secondary metabolites which apparently are produced by all
species of Allium. Firstly a range of sulphur compounds which originate all after wound-
ing from genuine S-alkyl- and S-alkenylcysteines and are responsible for the characteristic
odour. Secondly complex mixtures of biologically active saponins with C27 steroidal sapo-
genins. Thirdly phenolic compounds which seem to be mainly derivatives of the flavonols
Buijsen — Alliaceae 377
kaempferol and quercetin and simple phenolic acids such as protocatechuic and ferulic
acid. Perhaps biogenic amines and simple amides and alkaloids represent another group of
characteristic Allium metabolites.
The totality of the presently known chemical compounds indicates that Allium, and pos-
sibly Allioideae. have one taxon-characteristic chemical character, namely production of
S-alkylated cysteines. By their saponins Allioideae are reminiscent of that part of Lilii-
florae which is classified by Dahlgren et al. (1985) in Dioscoreales and Asparagales. Bio-
chemically Allioideae are clearly distinct from Amaryllidaceae s.str., Haemodoraceae,
Hypoxidaceae and all the families reunited in Liliales sensu Dahlgren et al. (1985).
Phytochemistry and chemotaxonomy of Alliaceae were discussed twice by Hegnauer
(1963, 1986) sub Liliaceae; in these treatises many references can be found. Some results
of recent phytochemical investigations will shortly be mentioned in the following alineas.
Sulphur compounds are usually considered to be mainly responsible for the medicinal
virtues of garlic and other species of Allium. This initiated a large number of chemical,
analytical and medicinal publications; see e.g. Ziegler et al. (1989), Sticher (1991), Block
(1992), Hikino et al. (1986) and others. The S-alkylated cysteines are stored in fresh bulbs,
leaves and seeds of Allium species as y-glutamyl peptides. During long storage or on
wounding a lot of mostly enzymatic transformations can take place, e.g.: generation of
the free S-alkylated cysteines; oxidation to S-alkylated cysteine sulfoxides (the S-allyl-
derivative is alliin); transformation of the sulfoxides to dialkyl thiosulfinates (the diallyl-
derivative is allicin); this last step is catalysed by the enzyme alliinase which is only
known from the genera Allium and Nothoscordum. S-rrarcs-l-propenylcysteine (the pre-
cursor of the lachrimatory factor of onion), S-allyl-cysteine (the precursor of alliin and
allicin), S-methylcysteine and S-propylcysteine occur in variable amounts and proportions
in different species of Allium (Lawson et ah 1991). New transformation products of
allicin are the ajoenes and the vinyldithiins of A. sativum (Sticher 1991). Onions produce
on grating and slicing variable amounts of thiopropanal-S -oxide (lachrimatory factor),
2-methyl-2-pentenal, propanethiol, dipropyldisulfide, propenyl-propyldisulfide and others
(Tokitomo & Kobayashi 1992). The cepaenes and deoxycepaenes of A. cepa are isomers
of the A. sativum ajoenes (Block & Zhao 1992). Other types of sulphur compounds were
isolated from subterranean parts ofTulbaghia violacea (Burton et al. 1992).
All species of Allium produce monodesmosidic spirostanol-type and bidesmosidic furo-
stanol-type steroidal saponins. A review treating 26 species, 26 sapogenins and 40 sapo-
nins was published by Kravets et al. (1990). Of the sapogenins mentioned agiogenin,
alliogenin, the ansurogenins, cepagenin, gantogenin, the karatavigenins, luvigenin, neo-
agigenin and its 6-benzoate and neoalligenin are new C27-spirostanols. Agiogenin was
first isolated from A. giganteum (therefore not aiogenin: Dahlgren et al. 1985: 195). New
saponins were isolated among others from bulbs of A. ampeloprasum (Morita et al. 1988),
bulbs of A. chinense (Matsuura et al. 1989a), bulbs of A. giganteum (Sashida et al. 1991),
flowers of A. porrum (Harmatha et al. 1987), bulbs and roots, but not leaves, of A. sati-
vum (Matsuura et al. 1988, 1989b) and bulbs of A. vineale (Chen & Snyder 1987, 1989).
The vineale saponins have molluscicidal activity; the leek saponin aginoside is concen-
trated in flowers and makes them unpalatable and toxic for larvae of the leek moth; the
spirostanolsaponins aginosideprosapogenin and ampeloside-Bsi of A. ampeloprasum are
378 Flora Malesiana ser. I, Vol. 11 (2) (1993)
fungitoxic whereas the bidesmosidic furostanol saponins ampeloside-Bf i and -Bf2 did not
inhibit the two species of Fungi tested; the same biological properties were observed in
A. sativum with the fungitoxic eruboside-B, a spirostanolic p-chlorogenin-3-glycoside,
and the inactive bidesmosidic furostanols proto-eruboside-B, sativoside-B 1 and sativo-
side-Rl. Bulbs of Tristagma uniflora yielded saponins with tigogenin, neotigogenin, two
25-epimeric 5a,6-dihydronuatigenins and two 25-epimeric 5a,6-dihydroisonuatigenins as
aglycones (Brunengo et al. 1985). According to Koch (1992) the steroidal sapogenins may
be involved in some of the therapeutical effects of onion and garlic.
Commercially available fresh leaves of A. tuberosum yielded three new kaempferol
bis- and tris-glycosides with one of the sugar hydroxyls acylated by ferulic acid, a kaemp-
ferol and a quercetin 3,4'-bisglucoside and kaempferol-3-sophoroside (Yoshida et al.
1987).
Bulbs of A. chinense (= A. bakeri) yielded diallyl disulfide, the dihydrostilbene lunu-
laric acid and the amides N-/?-coumaroyltyramine and N-feruloyltyramine (Okuyama et al.
1986; Goda et al. 1987). Another amide, aurantiamide acetate, was isolated from whole
plants of A. wallichii (Talapatra et al. 1989); these authors also reported isolation of the
furanocoumarin imperatorin from the same plant without giving yields nor mentioning
vouchers. A simple alkaloid related to N-methyltyramine and called alline (do not con-
found with alliin) was isolated from A. ramosum (= A. odorum) (Tashkhodzhaev et al.
1985) and sequently demonstrated to occur also in A. altaicum, anisopodium, senescens,
splendens, stellerianum and victorialis, but not in A. leucocephalum and A. schoenopra-
sum var. sibiricum (Antsupova & Polozhiy 1987).
Bulbs of all investigated Allium species store fructans (Hegnauer 1963; Deinko 1985)
and seeds store fatty oils with much linoleic acid (Hegnauer 1963; see for unsaturated fat-
ty acids of Allium taxa also Deinko 1985). According to Afzal et al. (1985) lipids of bulbs
of A. sativum contain much polyunsaturated fatty acids, such as linoleic, arachidonic
(= eicosa-all-ri.s-5,8,1 1,14-tetraenoic) and an eicosapentaenoic acid.
Phytoalexins were induced in bulb scales of A. cepa by Botrytis cinerea, and subsequent-
ly two fungistatic compounds could be isolated; they were called tsibulin-l,CnHi802,
and -2,Ci3H2202, after the Ukrainian name 'tsibulya' for onion; the tsibulins are 1-alkyl-
cyclopentan-2,4-diones (Tverskoy et al. 1991).
Van Damme et al. (1991) prepared lectins from bulbs of five species of Allium and
compared them with lectins of bulbs of six species of Amaryllidaceae s.str.; the taxon-
omic meaning of these results is not yet clear.
References: Afzal, M., et al., Agric. Biol. Chem. 49 (1985) 1187. — Antsupova, T.P. & A.V. Polo-
zhiy, Rast. Resur. 23 (1987) 436. — Block, E., Organosulphur chemistry of Allium, Angew. Chemie,
Intern. Ed. 31 (1992) 1135. — Block, E. & S.-H. Zhao, J. Org. Chem. 57 (1992) 5815. — Brunengo,
M.C., et al., Phytochemistry 24 (1985) 1388. — Burton, S.G., et al., Planta Medica 58 (1992) 295. —
Chen Shaoxing & J.K. Snyder, Tetrahedron Letters 28 (1987) 5603; J. Org. Chem. 54 (1989) 3679. —
Dahlgren, R.M.T., H.T. Clifford & P.F. Yeo. The families of the Monocotyledons (1985). — Deinko,
G.I., Rast. Resur. 21 (1985) 221 (lipids, fatty acids, carbohydrates). — Goda, Y., et al., Chem. Pharm.
Bull. 35 (1987) 2668. — Harmatha, J., et al., Biochem. Syst. Ecol. 15 (1987) 113. — Hegnauer, R.,
Chemotaxonomie der Pflanzen 2 (1963) 281,315-325,488-489,501; ibid. 7 (1986) 685-731,
803. — Hikino, H., et al., Planta Medica 52 (1986) 163 (antihepatotoxic action of garlic). — Koch, H.P.,
Zeitschr. Phytotherapie 13 (1992) 177 (ethnopharmacology of onion and garlic; modern pharmacological
Buijsen — Alliaceae 379
and clinical studies). — Kravets, S.D., et al., Khim. Prirod. Soedin. (1990) 429. — Lawson, D. A., et al.,
J. Nat. Prod. 34 (1991) 436. — Matsuura, H., et al., Chem. Pharm. Bull. 36 (1988) 3659; ibid. 37 (1989a)
1390 (chinensoside-I); ibid. 37 (1989b) 2741. — Merita, T., et al., Chem. Pharm. Bull. 36 (1988) 3480. —
Okuyama, T., et al., Planta Medica 52 (1986) 171. — Sashida, Y., et al., Chem. Pharm. Bull. 39 (1991)
698 (new alliogenin derivatives). — Sticher, O., Deutsch. Apoth. Z. 131 (1991) 403 (active principles
of garlic and their analytical control). — Talapatra, S.K., et al., Indian J. Chem. 28B (1989) 356. —
Tashkhodzhaev, B., et al., Khim. Prirod. Soedin. (1985) 687. — Tokitomo, Y. & A. Kobayashi, Biosci.
Biotechn. Biochem. 56 (1992) 1865. — Tverskoy, L., et al., Phytochemistry 30 (1991) 799. — Van
Damme, E.J.M., et al., Phytochemistry 30 (1991) 509. — Yoshida, T., et al., Chem. Pharm. Bull. 35
(1987) 97. — Ziegler, S.J., et al., Deutsch. Apoth. Z. 129 (1989) 318 (analytical control of garlic prepa-
rations).
R. Hegnauer
KEY TO THE GENERA IN MALESIA
la. Bruised leaves strongly smelling. Ovules 2 per locule (in the Malesian species); tepals
free or basally shortly connate Allium (p. 379)
b. Bruised leaves not or hardly smelling. Ovules many per locule; tepals basally shortly
connate Nothoscordum (p. 384)
ALLIUM
Allium L., Sp. PI. 1 (1753) 294. — Type species: Allium sativum L.
Herbs, usually with onion-smell, bulbs often present, sometimes with short rhizomes,
growing gregarious or not. Leaves linear to elliptic, sheathing the scape, the blades sub-
approximate or scattered along the scape, flat, or terete, semiterete, or angular and hollow.
Scape terete or angular. Inflorescence umbellate, usually many-flowered, sometimes with
bulbils, hemispherical to spherical or + ellipsoid, enveloped by 1 or 2 green or scarious,
persistent or caducous spathe(s). Flowers relatively small, white to pinkish or purplish,
bluish, or yellowish, stellate to campanulate or urceolate. Tepals 6, (sub)equal in length,
elliptic to lanceolate, free or usually shortly connate at base, persistent. Stamens 6; fila-
ments simple or tricuspidate. Ovary with 2-14 ovules per locule; style slender, erect;
stigma usually inconspicuous, single or 3-lobed. Fruit capsular, loculicidally 3-valved.
Seeds 1-4 per locule, flattened, irregularly angled, blackish.
Distribution — Indigenous to the Northern Hemisphere, with at present about 500-
700 species. In Malesia some taxa are cultivated, all are introduced, either from China,
Central Asia, or Europe. They are all well-known Allium crops, grown worldwide.
Uses — Of Allium plants the bulbs, cloves, pseudo-stems, leaves, and young inflores-
cences are used as vegetable and condiment. Allium species are also widely used in differ-
ent medicines for treating various diseases, e.g., diarrhoea, eye-infections, and head-
aches. Nowadays lowering of the blood pressure and inhibition of blood plate aggregation
are considered the most important medical effects of Allium consumption. Many Allium
species are grown as ornamentals; however, not so in Malesia because Allium species
usually do not flower under tropical conditions. For a more elaborate treatment of the cul-
tivated Allium species in SE Asia see Siemonsma & Kasem Piluek (eds.), PI. Res. SE
380 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Asia (PROSEA Haridb. 8, Vegetables) (1993). See also: Backer & Bakh.f., Fl. Java 3
(1968) 130-132; Burkill, Diet. Econ. Prod. Malay Penins. (1935) 98-103; Herklots,
Veget. SE Asia (1972) 387-401; Heyne, Nutt. PL Ned.-Indie, ed. 2 (1927) 439-442;
Jones & Mann, Onions and their allies. Botany, Cultivation, and Utilization (1963); Ochse
& Bakh., Veget. Dutch East Indies (1931) 440-457; Purseglove, Trop. Crops Monocot.
(1972) 37-57; Rabinowitch & Brewster (eds.), Onions and allied crops, Vol. 1 (1990).
KEY TO THE SPECIES CULTIVATED IN MALESIA
Where possible, vegetative characters are used, but sometimes additional generative characters are given.
Often young plants with immature bulbs are sold as vegetable. In that case characters of the bulb are not
fully reliable and other characters, e.g. leaves, should be checked.
1 a. Leaves terete or semiterete, or 3-5-angled, hollow 2
b. Leaves flat, sometimes V-shaped or keeled, not hollow 6
2a. Leaves terete or semiterete, with a wide central cavity, easily compressible, usually
diameter more than 5 mm (except A. schoenoprasum with a diameter of 1— 5(— 7)
mm) 3
b. Leaves 3- or 5-angled, with a narrow central cavity, diameter 1-5 mm
3. A. chinense
3a. Leaves terete, sometimes a few semiterete (check several leaves); mature bulbs dis-
tinct or indistinct, (narrowly) oblong to (narrowly) ovoid to elliptic 4
b. Leaves semiterete, flattened on upper surface (check several leaves); mature bulbs
distinct, depressed globose to ovoid or obovoid (immature bulbs of multiplier shallot
are narrowly oblong to narrowly ovoid). Variable! 2. A. cepa
4a. Bulbs indistinct, diameter of bulb and neck equal or diameter bulb up to 1.5 times
larger than diameter of neck 5
b. Bulbs distinct, diameter bulb 1.5-2 times larger than diameter neck
A. x proliferum Wakegi Group (see note under 4. A.fistulosum)
5a. Coarse plants; leaves fistulose, swollen in the centre, gradually tapering towards both
ends, usually diameter more than 5 mm; flowers pale yellow; stamens exceeding the
tepals 4. A. fistulosum
b. Plants of finer habit; leaves fistulose, not swollen, usually diameter less than 5 mm;
flowers usually purple, rarely white; stamens shorter than tepals
6. A. schoenoprasum
6a. Bulbs not set on a rhizome; leaves 20-60 cm by 10-35 mm 7
b. Bulbs with several closely together on a rhizome; leaves 13-45 cm by 2-10 mm
7. A. tuberosum
7a. Mature bulbs indistinct, ovoid to oblong, not composed of increase bulbs, increase
bulbs (if present) much smaller than main bulb 1. A. ampeloprasum
b. Mature bulbs distinct, depressed globose to ovoid, composed of cloves (increase
bulbs), cloves as long as main bulb (immature plants of A. sativum resemble A. am-
peloprasum but can be distinguished by the presence of cloves within the inner bulb-
coat-leaves and by a typical garlic-smell) 5. A. sativum
Buijsen — Alliaccac
381
1. Allium ampeloprasum L., Sp. PI. 1 (1753)
294.
Allium porrum L., Sp. PI. 1 (1753) 295.
Bulb indistinct, ovoid to oblong, diameter up to
c. 5 cm, gradually passing into the pseudo-stem.
Increase bulbs absent, few, or many, situated with-
in the outer bulbcoat-leaves. Foliage leaves 5-14,
suberect, upper part curved, flat, V-shaped in cross
section, up to 60 by 1-3.5 cm; sheaths extending
much above the ground, forming a pseudo-stem. In-
florescence (hemi) spherical. Flowers usually cam-
panulate, sometimes urceolate; tepals white to pur-
ple; stamens slightly shorter to longer than tepals.
Distribution - Allium ampeloprasum ranges as
a wild plant from S Europe and N Africa through
the Middle East into W and S Russia. A cultivated
form, also known as A. porrum, is grown mainly
in N Europe.
Common name - Leek (A. porrum).
2. Allium cepa L., Sp. PI. 1 (1753) 300.
Bulb distinct, depressed globose to ovoid or
obovoid, diameter 1— 10(— 15) cm. Increase bulbs
absent to several, ± ovoid or of similar shape as
the main bulb, often flattened on inner side. Pro-
tective bulbcoat-leaves several, papery, smooth,
purplish, yellowish, brownish or white; storage
leaves few to many. Foliage leaves 3-8(-9), erect
to suberect, semiterete, fistulose, 10-50 cm by
3-20 mm. Inflorescence spherical. Flowers sub-
campanulate to urceolate; tepals greenish to whitish;
stamens not or slightly exceeding tepals.
Two more or less distinct cultivar groups are
distinguished:
la. Bulbs small, diameter 1-3.5 cm; increase bulbs
several; protective bulbcoat-leaves purplish,
brownish, or white; plants 0.2-0.5 m, not
robust ... a. Cultivar group Aggregatum
b. Bulbs usually larger, diameter 2 — 10(— 15) cm;
increase bulbs absent or only few; protective
bulbcoat-leaves light yellow to light brown
or brown-purplish; plants 0.6-1.2 m, robust
b. Cultivar group Common Onion
a. Cultivar group Aggregatum
Allium ascalonicum auct. non L.: Ochse & Bakh.,
VegeL Dutch East Indies (1931) 441.
Allium cepa L. var. ascalonicum Backer, Handb.
Fl. Java 3 (1924) 60.
Bulb globose to ovoid, 1.5-4 by 1-3.5 cm.
Increase bulbs several, of similar shape as the
main bulb. Protective bulbcoat-leaves purplish,
brownish, or white. Foliage leaves 10— 35(— 4 1 )
cm by 3-10 mm.
Distribution - Centre of origin in N Africa and
E Mediterranean. Primary centre of diversity in the
Near East. Shallots are grown in the USA, Europe,
Africa, the Caribbean countries, Australia, and Asia.
Common names - Shallot, Multiplier shallot.
b. Cultivar group Common Onion
Allium cepa L. var. typicum Backer, Handb. Fl.
Java 3 (1924) 60.
Bulb depressed globose to ovoid or obovoid, 3-
5.5 by 2— 10(— 15) cm. Increase bulbs usually absent
or, in cultivars from India and Burma, few present,
± ovoid. Protective bulbcoat-leaves light yellow to
light brown or brown-purplish. Foliage leaves 35-
50 cm by 10-20 mm.
Distribution - The onion is not known as a wild
species, but in its primary centre of diversity, Cen-
tral Asia, several related wild species occur. A sec-
ondary centre of diversity is the Near East and the
Mediterranean, a rich diversity is also found in India
Common names - Onion, Dry bulb onion.
3. Allium chinense G. Don, Mem. Wem. Nat
Hist. Soc. 6 (1827) 83; Mann & Steam, Econ.
BoL 14 (1960) 69.
Allium exsertum (Lindley) Baker, J. Bot. (London)
12 (1874) 294, non G. Don (1827).
Allium bakeri Regel, Acta Horti Petrop. 3, iii
(1875)341.
Allium schoenoprasum auct. non L.: Heyne, NutL
PI. Ned. Indie' ed. 2 (1927) 441; Ochse & Bakh.,
VegeL Dutch East Indies (1931) 455; Backer &
Bakh.f., Fl. Java 3 (1968) 130.
Bulbs gregarious, narrowly ovoid, up to 3.5 cm
long, diameter 7-15 mm, gradually passing into
the leaves. Protective bulbcoat-leaves several, mem-
branous, smooth, white, brownish or purplish.
Foliage leaves 3-4(-5), prostrate to suberect, 3-
or 5-angled in section, 20-40 cm by 1-5 mm. In-
florescence umbellate. Flowers campanulate; tepals
light violet; stamens longly exceeding tepals.
Distribution - Native to C and E China. Culti-
vated in China, Japan, California, and SE Asia.
Carried worldwide as a garden crop by Asian com-
munities.
Note - Vegetative plants of A. chinense super-
ficially resemble those of A. schoenoprasum. How-
ever, the latter species has terete leaves and indis-
tinct bulbs, whereas A. chinense has more or less
angled leaves and distinct bulbs.
Common name - Rakkyo.
382
Flora Malesiana ser. I, Vol. 11 (2) (1993)
t
Fig. 1. Allium fistulosum L. a. Habit; b, c. outer and inner tepal with corresponding filaments. From:
B.E.E. de Wilde-Duyfjes, A revision of the genus Allium in Africa [Meded. Landbouwhogeschool
Wageningen 76-11 (1976) 88, fig. 15].
Buijsen — Alliaceae
383
4. Allium fistulosum L., Sp. PI. 1 (1753) 301.
Bulb indistinct, oblong to ovoid, 3.5-8 cm
long, width 5-25 mm, gradually passing into the
leaves. Increase bulbs few to several, narrow and
inconspicuous. Protective bulbcoat-leaves several,
papery, smooth, brownish, whitish, or purplish.
Foliage leaves 4-6, erect, terete, fistulose, 20-54
cm by 5-20 mm. Inflorescence (hemi)spherical.
Flowers narrowly campanulate to urceolate; tepals
pale yellow; stamens longly exceeding tepals. -
Fig. 1.
Distribution - Origin in Siberia and China, not
known as a wild species. Allium fistulosum has
been, since prehistoric times, the main garden
onion of China and Japan. It is cultivated widely
throughout the world, ranging from Siberia to trop-
ical Asia, including China, Japan, Korea, Taiwan,
and the SE Asian countries.
Common names - Japanese bunching onion,
Welsh onion, Green bunching onion.
Note - Allium x proliferum (Moench) Schrad.
ex Willd. Wakegi group, a hybrid between A. fis-
tulosum and A. cepa cv. group Aggregatum, is
cultivated occasionally in Malesia. It has been
grown for centuries in China, Japan, and SE Asia
[Hanelt in Rabinowitch & Brewster (eds.), Onions
and allied crops 1 (1990) 18]. This hybrid has char-
acters from both parents. It develops a distinct bulb
like the shallot, and terete leaves like A. fistulo-
sum but slender and very erect. The flowers are
subcampanulate as in A. cepa, but the stamens ex-
ceed the perianth and the inner filaments are with-
out lateral teeth at the base as in the other parent.
The hybrid is completely sterile and does not form
ferule seeds.
5. Allium sativum L., Sp. PI. 1 (1753) 297.
Bulb distinct, depressed globose to ovoid, dia-
meter up to c. 7 cm, mainly composed of increase
bulbs (cloves). Increase bulbs (l-)4-15, broadly
ovoid to ovoid, as long as the main bulb when
mature, wrapped in one cartilaginous prophyll.
Protective bulbcoat-leaves papery or chartaceous,
smooth, whitish or purplish, in young plants
fibrous and brown. Foliage leaves 4-\0, curved,
flat, V-shaped in section, 20-50 cm by 10-25
mm. Inflorescence subspherical, composed either
of only sessile bulbils (topsets) or of bulbils mixed
with flowers. Flowers usually ill : devcloped, or
rudimentary, or absent; perianth subcampanulate;
tepals pale pink or greenish; stamens shorter than
tepals.
Distribution - Some consider A. longicuspis
Regel, endemic to Central Asia, as the wild parent
of this cultigen. It was originally grown in Europe
and China and nowadays worldwide.
Common name - Garlic.
6. Allium schoenoprasum L., Sp. PI. 1 (1753)
301.
Bulbs gregarious, indistinct, oblong to narrow-
ly ovoid, 1-3 cm long, gradually passing into scape
and leaves, several set on a rhizome. Increase bulbs
few to several, narrow and inconspicuous. Protective
bulbcoat-leaves several, papery, smooth, brownish.
Foliage leaves 3-6, erect, terete, fistulose, 10-50
cm by 1— 5(— 7) mm. Inflorescence (hemi)spherical.
Flowers narrowly urceolate; tepals white to purple;
stamens much shorter than tepals.
Distribution - This very variable species is
known as a wild plant throughout the Northern
Hemisphere: Europe, Asia, and North America. It
is cultivated worldwide, in Malesia it is grown only
occasionally.
Common name - Chive.
Note - Allium schoenoprasum is sometimes
confused with A. chinense, see under the latter spe-
cies.
7. Allium tuberosum Rottier ex Spreng., Caroli
Linnaei SysL Veg. 2 (1825) 38; Steam, Her-
bertiall (1944)226.
Allium uliginosum G. Don, Mem. Wem. Nat.
Hist. Soc. 6 (1827) 60.
Allium odorum auct. non L.: Heyne, Nutt. PI. Ned.
Indie ed. 2 (1927) 441; Ochse & Bakh., VegeL
Dutch East Indies (1931) 450; Backer & Bakh. f.,
Fl. Java 3 (1968) 131.
Allium bakeri aucL non Regel: Backer & Bakh. f.,
Fl. Java 3 (1968) 131.
Bulb indistinct, narrowly ovoid to ovoid, 15-20
by c. 15 mm, several set on a rhizome. Protective
bulbcoat-leaves several, broken up into netted fibres,
light brown to brown. Foliage leaves 4-9, sub-
erect or curved, flat, slightly rounded or keeled on
lower surface, 13-45 cm by 2-10 mm. Inflores-
cence umbellate. Flowers stellate; tepals white;
stamens ± equalling tepals.
Distribution - The primary centre of origin is
unknown, as this species easily runs wild. Known
as a wild species in eastern Asia. Cultivation from
E Mongolia to Japan, the Philippines, Indonesia,
Malaysia and through Thailand to N India.
Common name - Chinese chive.
Note - Sometimes sub-naturalized.
384
Flora Malesiana ser. I, Vol. 11 (2) (1993)
NOTHOSCORDUM
Nothoscordum Kunth, Enum. PL 4 (1843) 457, nom. gen. cons.
Nothoscordum inodorum (Aiton) G. Nichols,
111. Diet. Gard. 2 (1885/89) 457; Backer &
Bakh. f., Fl. Java 3 (1968) 132; Jessop in Fl.
Males. I, 9 1 (1979) 234.
Glabrous, inodorous herb with bulbs. Leaves
radical, linear, flat, 15-45 cm by 5-12 mm. Um-
bels on up to 70 cm long peduncle, 6-17-flowered.
Tepals 6, shortly connate at base, white, some-
times with a purple median streak. Stamens 6, in-
serted on the base of the perianth; filaments ligu-
late. Ovary with many ovules per locule. Fruit a
capsule, loculicidally 3-valved. Seeds several, black.
Distribution - Native to subtropical North
America; cultivated as an ornamental, often escaped
and naturalized. In Java sometimes cultivated,
locally naturalized in W Java (Backer & Bakh. f.,
I.e.).
Habitat & Ecology - Sometimes gregarious in
fields, tea-gardens, and along roadsides, altitude
1000-1500 m. Fl.: Jan.-Dec. Easily propagated
by bulbils and seeds, difficult to eradicate.
Note - The species was mentioned by Jessop
(I.e.) in the Flora Malesiana treatment of Lilia-
ceae, in which treatment the cultivated vegetables
and condiments were ignored.
Flora Malesiana ser. I, Vol. 11 (2) (1993) 385-391
CORIARIACEAE
(Brigitta E. E. Duyfjes, Leiden, The Netherlands)
CORIARIA
Coriaria L., Sp. PI. ed. 1 (1753) 1037. — Lectotype species: Coriaria myrtifolia L.
Shrubs (rarely perennial herbs with herbaceous sterns) or small trees. Roots with nodules
with nitrogen fixing bacteria. Innovations in flushes from perulate apical buds. Branches
4-angular, ridges alternating; xylem with broad medullary rays. Leaves generally small,
sessile or subsessile, with a narrow joint at base, simple, entire, cordate-ovate to lanceo-
late, acute, decussate, or (not in Malesia) in whorls, phyllotaxis of the lateral branches
mostly showing as pseudo-distichous; blades palmately 3-9-curvi-nerved, net-veined,
those sustaining side-branches often of somewhat different shape. Stipules absent (but see
note). Racemes terminal or axillary. Rowers in the axils of small bracts, greenish or red-
dish, actinomorphic, hypogynous, polygamous, i.e., bisexual or functionally unisexual
(both male and female flowers with rudiments of the other sex). Pedicel slender, brac-
teoles absent. Perianth 5-merous; sepals 5, imbricate, persistent, ovate; petals 5, alternate,
shorter than the sepals, keeled inside, accrescent, becoming more or less fleshy, each
enveloping a carpel in fruit. Stamens 10, in two whorls, free, or those opposite the petals
adnate to the keel of the petal; filaments filiform; anthers exserted, ellipsoid or sagittate,
papillose or smooth, 2-celled, opening lengthwise. Carpels 5— 10(— 12), free, one-celled,
the basal part adnate to a central conical receptacle, each with a slender distinct papillose
style. Ovules one per carpel, pendulous, anatropous, with the micropyle directed upwards.
Pseudo-fruit consisting of hard-walled, laterally compressed achenes, more or less en-
closed by the fleshy-accrescent petals, each achene subtended on each side by half of the
adjoining petal. Seed compressed, without an aril, testa thin, endosperm scanty or none;
embryo straight. — Fig. 1.
Distribution — About 20 species, depending on the taxonomic treatment (see infra-
generic variation), with a much broken distribution: the West-Mediterranean, the Hima-
layan Region from Pakistan to SE Asia, and E Asia (Japan, China, incl. Hainan), through
Malesia (rare) to New Zealand (and the Subantarctic Islands), Chile, Peru as far North as
Mexico; in Malesia: the Philippines (Luzon) and Papua New Guinea.
Habitat & Ecology — In Malesia and SE Asia in the submontane and montane zone,
often pioneering in exposed situations. Apparently a genus of temperate and subtropical
climatic conditions. There are remarkable disjunctions in the distribution. The flowers,
with exerted slender stamens and styles, are adapted to wind pollination. The fleshy peri-
carp suggests dispersal by animals, presumably birds. The racemes and fruits have a
superficial resemblance with the Phytolaccaceae.
(385)
386 Flora M alesiana ser. I, Vol. 11 (2) (1993)
Systematic position of the family — This monogeneric family has been the subject of
various studies in an attempt to settle its phylogenetic and systematic position. It has no
close relationship to any suggested family. Its distribution and the numerous proposed
putative alliances give the impression of an old genus; it is also found as a fossil (see the
chapter on palynology). Possibly the status of a separate order Coriariales for this single
isolated genus is a good solution.
References: Bohm, B.A. & R. Ornduff, Syst. Bot. 6 (1981) 15-26. — Carlquist, S., Syst. Bot. 10
(1985) 174-183. — Cronquist, A., An integrated system of classification of flowering plants (1981) 136-
139; The evolution and classification of flowering plants (1988) 289, 504. — Engler, A., in Engler &
Prantl, Nat. Pflanzenfam. Ill, 5 (1896) 128-129. — Garg, M, Phytomorphology 30 (1980) 15-26. —
Hutchinson, J., Families of flowering plants (1926) 156; ibid., ed. 3 (1973) 136, 183-184. — Oginuma, K.,
M. Nakata, M. Suzuki, & H. Tobe, Bot. Mag. Tokyo 104 (1991) 297-308. — Takhtajan, A., Evolution
und Ausbreitung der Bliitenpflanzen (1973) 173; Bot. Rev. 46 (1980) 225-259; Floristic regions of the
world (1988) 334.
Infrageneric variation — Because of the great variation within populations of Coriaria,
the generally weak demarcation of species described and the occurrence of natural hybridi-
zation, especially apparent in New Zealand, Skog (1972) distinguished only five species
in total for the whole genus. One of these is the aggregate species C. ruscifolia L., com-
prising all species from South and Central America, the South Pacific islands, New Zea-
land and New Guinea. Coriaria ruscifolia has two subspecies, subsp. ruscifolia and subsp.
microphylla, the latter including C. papuana from New Guinea. Following Skog, our
present C. intermedia would be included in C.japonica. To the contrary, a recent karyo-
morphological study (Oginuma et al. 1991) provides evidence for distinguishing at least
four species in C. ruscifolia sensu Skog. For convenience sake, two locally well-charac-
terized species are accepted here for Malesia, C. intermedia and C. papuana.
References: Oginuma, K., M. Nakata., M. Suzuki & H. Tobe, Bot. Mag. Tokyo 104 (1991) 297-
308. — Skog, L.E., Rhodora 74 (1972) 242-253.
Vegetative Anatomy — Leaf anatomy. Cuticle striated; stomata paracytic; lamina dor-
siventral; midrib with single collateral vascular strand supported by weakly developed
sclerenchyma caps.
Wood anatomy. Wood weakly semi-ring-porous or diffuse-porous. Vessels angular,
thin-walled, in multiples and small clusters, rarely solitary, with simple perforations and
alternate intervessel pits with slit-like, occasionally coalescent apertures. Vascular tracheids,
resembling narrow vessel elements, present in association with vessel groups. Ground
tissue composed of libriform fibres. Parenchyma vasicentric to confluent, fusiform or in
2(-4)-celled strands. Rays multiseriate, broad and tall, composed predominantly of erect
and square cells, integrading with procumbent cells. Crystals present in some ray cells.
Vessel elements, axial parenchyma, and sometimes also the fibres storied.
Carlquist (1985) concluded that the affinities of Coriaria remain elusive, despite some
wood anatomical similarities with Corynocarpus, woody Ranunculaceae, and several
Simaroubaceae.
Duyfjes — Coriariaceae 387
References: Carlquist, S., Syst. Bot. 10 (1985) 174-183 (wood anatomy of Coriaria arborea, C.ja-
ponica, C. ruscifolia and C. thymifolia) — Metcalfe, C.R. & L. Chalk, Anatomy of the Dicotyledons
(1950) — Santos, J.K., Philipp. J. Sc. 46 (1931) 257-266 (leaf anatomy of Coriaria intermedia). —
Suzuki, M. & K. Yoda, J. Jap. Bot. 61 (1986) 289-296, 333-341 (wood anatomy of Coriaria termi-
nalis, C. japonica, C. nepalensis and C. intermedia). — Yoda, K. & M. Suzuki, Bot. Mag. Tokyo 105
(1992) 235-245 (wood anatomy of 14 species, including also Coriaria intermedia).
P. Baas
Chromosomes — The basic chromosome number is x = 20; the somatic chromosome
number is for the majority of species, including the Malesian representatives, 2n = 40, but
outside Malesia tetraploidy (2n = 80) also occurs.
Reference: Oginuma, K., M. Nakata, M. Suzuki, & H. Tobe, Bot. Mag. Tokyo 104 (1991) 297-308.
Palynology — Coriaria pollen is small to medium-sized (polar axis 14-35 u.m, equa-
torial diameter 20-34 urn), isopolar, usually suboblate (P/E 0.75-0.88) and 3-aperturate,
either colporate or pororate. The equatorial outline is (sub)circular, the meridional outline
elliptic. The colpori have an ectocolpus of 4-10 by 0.5-2 u.m and a lalongate eridoporus
of 2-5 by 3-8 (im. Pororate grains have isodiametric ectopori of 2-4 jim, often with
thickened protruding margins, and lalongate to isodiametric endopori of 3-5 by 4-6 u.m.
The exine is 0.75-1.5 u.m thick and distinctly stratified, showing a nexine columellate
infratectal layer and a tectum, which are about equally thick. The tectum has a scabrate or-
namentation consisting of c. 0.2 |im high conical elements and shows many small funnel-
shaped perforations. Individual columellae, scabrae and perforations are usually not dis-
tinguishable with light microscopy.
Praglowski (1970) studied the pollen of 16 species. He recognised a group with col-
porate pollen (12 species from America, the Mediterranean region, China, New Guinea
and New Zealand) and a group with pororate pollen (4 species from India, China, Japan,
Taiwan and Luzon). The colporate C. terminalis from China is intermediate, having rather
short ectocolpi (3-5 u.m).
Skog (1972) distinguished only 5 species in Coriaria: C. ruscifolia (including all mate-
rial from America, New Guinea, New Zealand and other Pacific islands) has colporate
pollen; C. terminalis (Tibet, China) and C. myrtifolia (Mediterranean region) are two other
colporate species; C. nepalensis (including C. sinica; India, Nepal, China) and C. japoni-
ca (including C. intermedia; Japan, Taiwan, Luzon) have pororate pollen.
Pollen morphology does not elucidate the phylogenetic position of Coriaria. Coriaria
pollen has been tentatively compared with that of Sapindaceae (Meenakshi Garg 1980),
but obviously the available data of the latter did not represent adequate reference material.
Coriaria pollen is relatively 'simple', and does not show special features that allow affilia-
tion with other taxa. It resembles pollen of Corylus (Betulaceae) and Myrica (Myricaceae).
However, this resemblance may be explained from the anemophilous pollination system
shared by these plants.
Fossil Coriaria pollen has been reported from the upper Miocene of Spain, the Pliocene
of New Zealand (see Muller 1981) and the Pleistocene of the Netherlands (see Praglowski,
I.e.).
388 Flora Malesiana ser. I, Vol. 11 (2) (1993)
References: Meenakshi Garg, Phytomorphology 30 (1980) 5-10. — Muller, J., Bot. Review 47
(1981) 1-142. — Praglowski, J., World Pollen Flora 1 (1970) 15-22, 25-31. — Skog, L.E., Rhodora
74 (1972) 242-253.
R.W.J.M. VanDerHam
Phytochemistry — The monogeneric family has its name from the Latin corium
(= leather) because leaves and twigs of the Mediterranean Coriaria myrtifolia were former-
ly used in tannery. Chemical characters of Coriariaceae were summarized by Hegnauer
(1964, 1989), where many references to phytochemical and toxicological literature are
available. The phytochemistry of this small family is surprisingly well known. All species
seem to contain bitter, toxic lactones and to produce large amounts of ellagitannins accom-
panied by lesser amounts of precursor gallotannins.
The toxic, lactonic principles are picrotaxan-type sesquiterpenes. Coriamyrtin, corianin,
tutin (from the Maori name 'Tutu' of the genus), and pseudotutin are the toxic picrotoxin-
like compounds of Coriaria. All Coriaria species are known to be toxic. The picrotaxans
are a group of convulsive, insecticidal and ichthyotoxic natural products which occur erra-
tically in angiosperms (also known from a few genera of Menispermaceae, Euphorbiaceae,
Orchidaceae).
The ellagitannins of Coriaria japonica were studied thoroughly by Okuda's group
(Okuda et al. 1990, 1993). Monomeric ellagitannins like tellimagrandin-I and -II, coriariin-B
and -F and geraniin and dimeric ellagitannins like coriariin-A and -C to -E, and rugosin-D
and -E are produced by this taxon which also contains trigalloylglucose.
Glycosides of the flavonols kaempferol and quercetin are present in leaves of all species
which have been investigated. They are accompanied in most species by the 7-glucoside
of the flavanone naringenin.
Fruits, leaves and twigs of the South American species C. ruscifolia yielded the triter-
penic acid ursolic acid and phytosterins and coriamyrtin.
The carbohydrate metabolism of the family shows special features. A new heptulose,
coriose, was isolated from roots, stems, leaves, fruits and seeds of C. japonica; it is ac-
companied by sedoheptulose in leaves and by a corresponding heptitol, volemitol, in
seeds. Coriose and another heptulose are also present in C. intermedia, nepalensis, rusci-
folia and thymifolia. A/yo-inositol is also present in easily detectable amounts in all parts
of C. japonica.
Seeds contain much fatty oil (up to 70% of kernels) with an unusual fatty acid, coriolic
acid, as the main fatty acid. Coriolic acid is 9-cis,\ l-fr<ms-13-hydroxyoctadecadienoic acid;
this acid can be interpreted as a monohydrated a-elaeostearic acid. Seed oils of all investi-
gated species, C. myrtifolia, nepalensis, papuana, ruscifolia, sarmentosa and terminalis,
contained this new fatty acid.
On account of its chemistry the family must belong to the core of dicotyledons which
produce and accumulate ellagitannins.
Phytochemistry suggests Euphorbiaceae on the one side and Crassulaceae-Saxifraga-
ceae s.str. on the other side as good candidates for remote relatives. Thome (1992) clas-
sifies Coriariaceae in Rutanae in the most recent version of his phylogenetic system of
angiosperms and Huber (1991) suggests close relationship with Anacardiaceae.
Duyfjes — Coriariaceae 389
References: Huber, H., Angiospermen. Leilfaden durch die Ordnungen und Familien der Bcdecktsamer
(1991), G. Fischer, Stuttgart. — Okuda, T., et al., Heterocycles 30 (1990) 1195 (review of oligomeric
hydrolisable tannins; a new class of plant polyphenols). — Okuda, T., et al., Phytochemistry 32 (1993)
507 (classification and distribution in dicotyledons of oligomeric hydrolisable tannins). — Thome, R.F.,
Classification and geography of the flowering plants, Bot. Rev. 58 (1992) 225-348.
R. Hegnauer
U ses — Rich in tannin, used locally for tanning and black-dye. The flowers and fruits
contain a narcotic, poisonous substance, coriariin. Some Coriaria species (e.g. the Medi-
terranean C. myrtifolia L., some New Zealand species, and also C. papuana) possess root
nodules with nitrogen-fixing properties. As these species often grow in pioneering situa-
tions, such as lava fields and roadsides, they may have a place in planting programmes for
erosion control in wet scree country (Daly et al. 1972). Coriaria nepalensis (India) is one
of the food plants of the silk moth.
References: Ambasta (ed.), Useful plants of India (1986) 142. — Daly, G.T., B.E. Smith & S. Chua,
Proc. New Zeal. Ecol. Soc. 19 (1972) 65-74. — Watt, G., The commercial products of India (1908) 1012.
Note — In and beside the leaf-axils numerous minute, fleshy, finger-shaped, and gland-
like emergentia are usually present. These have been called stipules by various authors.
KEY TO THE SPECIES
la. Racemes 4-10 cm long, axillary, aggregated in groups of (1 or) 2 or 3, subtended by
small cataphylls. Flowers c. 5 mm across. Leaves oblong, 5-7 cm long, 3-nerved.
Fruits c. 5 mm across 1. C. intermedia
b. Racemes (7-)20 cm long, axillary, solitary. Flowers c. 2.5 mm across. Leaves o r ate-
oblong, 1.5-4 cm long, 5 -7 (-11)- nerved. Fruits 2-3 mm across 2. C. papuina
1. Coriaria intermedia Matsumura, BoL Mag. broadly ovate, c. 3.2 by 2.4 mm, obtuse; pelvis
Tokyo, Bot. Soc. 12 (1898) 62 ; Hui-Lin Li, ovate, c. 1.2 by 0.5 mm, acute; stamens c. 8 mm
Woody Fl. Taiwan (1963) 443; in Fl. Taiwan 3 long, filaments c. 6 mm long, anUvs 1.5-2 by
(1977) 563, pi. 729; Kanehira, Formos. Trees, c. 0.8 mm, papillose. Female flowers: sepals
rev. ed. (1936) 361, f. 317; Keng, Orders and broadly ovate, c. 3 by 2 mm, obtuse, or acute;
families of seedplants of Taiwan (1987) 62, f. petals ovate, c. 2.8 by 1.8 mm, acute; styles c.
20.1; Merr., Enum. Philipp. Flow. PI. 4 3.5 mm long; carpels 5, c. 1.5 by 1 mm. Pseudo-
(1926) 88. — Types: Tashiro s.n. (not seen), fruit 3-5 mm across; achenes 2.8-3 by 1.5-1.8
Owatari s.n. (not seen), both from Taiwan. mm, glabrous, with a few longitudinal ribs.
Shrubs or small trees, 1-3 m high. Roots form- Distribution - Taiwan; in Malesia: Philippines
ing nodules. Leaves decussate; petioles c. 2 mm ^ N Luzon).
long; blades oblong (or ovate), glabrous, 3-nerved, Habitat & Ecology - Thickets, open forests,
midrib pinkish, (3-)5-7 by 1.5-2.5 cm, base sunny and stony slopes, dry riverbeds and water-
cuneate, apex acute. Inflorescences consisting of courses, from low altitudes up to c. 2400 m.
subaxillary fascicles of 1-3 racemes, 4-10 cm Notes - Closely allied to C. japonica.
long, glabrous; bracts ovate, glabrous, 4.5-5 by Due to its occurrence in riverbeds and water-
2.5-4 mm, acute or blunt, the margins irregularly courses, C. intermedia is possibly a facultative
serrate. Flowers bisexual or unisexual, yellowish rheophyte, but it was not recognized as such by
green; pedicels 2-7 mm long. Male flowers: sepals Van Steenis (1981); see also under C. papuana.
390
Flora Malesiana ser. I, Vol. 1 1 (2) (1993)
Fig. 1. Coriaria papuana Warburg, a. Portion of branch and apical part of branch with inflorescences,
x 0.6; b, c. flowers, young and in anthesis, x 6; d. perianth, stamens and pistil removed, x 6; e, f.
immature and mature stamens, x 10; g. pistil, x 10 (drawn by R. van Crevel after a specimen collected by
Gyldenstolpe in herb. S).
Duyfjes — Coriariaceae
391
According to Hui-Lin Li (1963, 1977) the roots
and seeds are very poisonous. Keng (1987) men-
tioned that the species is used in local medicine.
Bisexual and unisexual flowers: in the studied
material, besides bisexual flowers with normal
looking stamens and pistils, only unisexual male
flowers were present, i.e. flowers with well-devel-
oped stamens, destitute of pistillodes; unisexual
female flowers have never been seen. Most flowers
had already developed pseudo-fruits with persisting
withered stamens.
2. Coriaria papuana Warb., Bol Jahrb. 16
(1893) 22, f. 741; P. van Royen, Alpine Flora
New Guinea 4 (1983) 2547; Steenis, Rheophytes
of the World (1981) 219. — Type: Hellwig 364
(B, lost); neotype: van Royen 16242 (L).
Spreading and much branched bushy shrubs or
small trees 0.5-4 m high; young flushes tinged
reddish. Root system extensive and forming nod-
ules. Leaves decussate, subsessile, ovate-oblong,
base rounded to cordate, apex acute; (l-)1.5-4 by
0.8-1.7 cm, glabrous, except nerves on both sur-
faces with short white hairs; nerves 3-5; leaves at
the base of lateral twigs and inflorescences smaller,
cordate, up to 1 1 -nerved, inflorescences consisting
of solitary axillary racemes 7-20 cm long, rhachis
and pedicels densely beset with short white hairs;
bracts ovate to lanceolate, 2-3 mm long, shortly
acuminate, glabrous or scabrous on the lower sur-
face, margin irregularly serrate. Flowers bisexual,
dark red, pink or yellow-green; pedicels 3-7 mm
long; sepals broadly ovate, 1.5-2 by 1-1.2 mm.
acute; petals broadly ovate, 1-1.8 by 1-1.5 mm,
acute. Stamens 2-3 mm long; filaments filiform,
0.5-1 mm long; anthers ellipsoid, 1-1.2 mm long,
papillose. Carpels 5, c. 0.8 by 0.4-0.8 mm; styles
c. 3 mm long. Pseudo-fruits 2-3 mm across, red,
dark purple or blackish, soft when ripe; achenes
c. 1.5 by 1.2 mm, glabrous, with a few longitudi-
nal vein-like ribs. - Fig. 1.
Distribution - Malesia: endemic to Papua New
Guinea: Western Highlands (Chimbu), Eastern
Highlands, Morobe Prov. (Finisterre Mts, Mt Dick-
son area).
Habitat & Ecology - Alongside and in stony
and rocky riverbeds, on foothills and in upper mon-
tane forest; altitude 600-3500 m. Frequently grow-
ing gregariously as a pioneer in exposed places
like ridges and landslides.
Note - Because of the enormous root-system
and the preference for growing in riverbeds, the
species was listed as a facultative rheophyte by
Van Steenis (1981).
Flora Malesiana ser. I, Vol. 1 1 (2) (1993) 393-398
PENTASTEMONACEAE
(Brigiua E. E. Duyfjes, Leiden, The Netherlands)
The taxonomic position and rank of the only genus Pentastemona has been under discus-
sion since Van Steenis (1982) described the genus in the Stemonaceae. Dahlgren et al.
(1985) found it highly distinctive and suggested it worthy of family rank. Later on more
material off. sumatrana and P. egregia has become available facilitating a more accurate
description of the androgynoecium. Conspicuous differences from the Stemonaceae, viz.
the inferior ovary, the five pouches caused by the fusion of connectives and stigma (espe-
cially well-developed in P. egregia), the berry-like fruit and the exotesta (sarcotesta) of the
seed, warranted the distinction of a separate new family for the genus.
Note — Pentastemona is the first genus in the Monocotyledons with normally regular
5-merous flowers. Checked on abundant material, occasional flowers with 4 or 6 perianth
lobes and stamens occur in both species.
References: Dahlgren, R.M.T., H.T. Clifford & P.F. Yeo, The families of the Monocotyledons
(1985) 123. — Duyfjes, B.E.E., Blumea 36 (1991) 239-252; ibid., 36 (1992) 551-552 (family de-
scription). — Steenis, C.G.GJ. van, Blumea 28 (1982) 151-163, f. 1.
PENTASTEMONA
Pentastemona Steenis, Blumea 28 (1982) 160. — Type: Pentastemona sumatrana Steenis.
Low juicy herbs with a short, unbranched, largely overground green rhizome with pale
roots; stem curving up into a short erect portion bearing several leaves, leaving thin, al-
most ring-shaped scars; plants almost glabrous or with sparse uniseriate hairs. Leaves
papyraceous when dry; blade ovate, with 6-8 pairs of arching basal and suprabasal
nerves, shallowly depressed above, secondary intervenation fine-trabeculate, margin
entire; petiole long, towards the base canaliculate and shortly sheathing, with the edge
hyaline and fringed- hairy. Inflorescences axillary, either short-peduncled simple racemes
or long-peduncled compound racemes; flowers and bracts dotted with raphides, bracteoles
absent. Flowers with one kind of perianth, 5-merous; tepals free or partly united, imbri-
cate, outside papillose, persisting in fruit. Stamens epitepalous; filaments apparently ab-
sent since they are completely united into a conspicuous fleshy ring, which is either free
or partly fused to the base of the fleshy flower tube, which, together with the juicy-fleshy
torus and the stigma, form a solid hypanthium with a flat or concave top; anthers, consist-
ing of two short, broadly ovate thecae, each opening by a longitudinal lateral slit. The
thecae are sessile, laterally directed and separated by broad fleshy connectives, which
taper into slender or broad inward-curved appendices, the tips of which are fused with the
stigma; the stamens and stigma thus form 5 pouches in the fleshy hypanthium, each con-
taining two thecae, one from each of the adjacent anthers. Viewed from above, the ap-
pearance is as if the anthers are alternitepalous (P. egregia), or anthers shortly protruding
(393)
394
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 1. Pentastemona sumatrana Steenis. a. Habit, x 0.5; b. base of leaf sheath, x 1.5; c. flower, x 6;
d. flower, tepals removed, x 6; e. ribbed seed, sarcotesta torn away, x 25. — P. egregia (Schott) Steenis.
f. Rower, x 2 (a-e: de Wilde & de Wilde-Duyfjes 18695; f: Meijer 17010).
Duyfjes — Pentastemonaceae 395
(P. sumatrana). Ovary inferior, one-celled, ovules numerous on three inward projecting
longitudinally furrowed, parietal placentas; style short and inconspicuous, stigma broad,
flattish, entire or 3- or 4-lobed, papillose. Fruit berry-like, sharply longitudinally 10-ridged,
the perianth persisting. Seeds many, with a collar-like undulate aril covering about one
third of the seed; exotesta (sarcotesta) transparent; pericarp strongly ribbed; endosperm
large; embryo small. — Figs. 1-3.
Distribution — Malesia: the genus is locally endemic to N and W Central Sumatra,
with 2 species.
Habitat & Ecology — Gregarious but very local. In damp places on rocks in rain for-
est of lowland and hills and apparently often more or less kremnophytic. The berry-like
fruit probably decays, by which the seeds become free. When a mature fruit of P. suma-
trana is opened, the seeds, with their ribbed endocarps, lie in a jelly-like mass of the arils
and exotestas, appearing as one coherent mass and suggesting an as yet unknown mode
of dispersal.
Pentastemona sumatrana may produce young plants at the top of the inflorescences in
a viviparous manner, while in P. egregia young plants easily sprout from the margins of
broken leaves in their natural habitat. Under damp hothouse conditions P. egregia can be
propagated in this way.
There are indications that the family possibly is dioecious. The collection de Wilde
20113 of P. sumatrana contains solely female flowers and fruits with ripe seeds; the an-
thers, although present, appeared to be empty. The collection Bogner 1724 of P. egregia
bears functionally male flowers, the anthers of which contain good pollen. Ovaries are
also present in these flowers, but are not well-developed and probably contain abortive
ovules.
According to Van Steenis (1982) the smallish deep pouches in the receptacle, as in
P. egregia, suggest nectarial structures, but microscopical examination proved that such
structures are absent.
Although gregarious in their natural habitat, the plants themselves are not conspicuous
and may have escaped the attention of collectors. This may be a reason for their late dis-
covery.
Floral morphology — The androecium shows a corona-like development of the basal
staminal region like that in Peliosanthes {LiliaceaelConvallariaceae). The late differentia-
tion of the stamens into extensions that contact the stigma, as well as the inferior position
of the ovary, suggest a relationship with Trichopus (Trichopodaceae) and Stenomeris
(Stenomeridaceae).
Reference: Heel, W. A. van, Blumea 36 (1992) 481-499, pi. 6, 7.
Seed — The seed shows a remarkable character, viz., a distinct, proportionally thick
exotesta (sarcotesta) which in dried condition is only little visible. For the description of
ovules and seeds, see Bouman & Devente (1992).
Reference: Bouman, F. & N. Devente, Blumea 36 (1992) 501-514, pi. 1, 2.
396
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 2. Pentastemona, lateral views of mature flowers, a, b. P. sumatrana Steenis, male and female flower
respectively; c, d. P. egregia (Schott) Steenis, ditto; all x 4 (a, b: de Wilde & de Wilde-Duyfjes 21399;
c: Bogner 1724; d: Meijer 17010).
Anatomy — Leaf epidermis anticlinal cell walls straight; stomatal complex tetracytic to
cyclocytic; crystals present as abundant styloids and raphides, also as minute, irregularly
shaped, solitary crystals. Minute druses have been found throughout the mesophyl. For
comparison with Stemonaceae, see there (p. 400).
Reference: Baas, P., in C.G.G.J. van Steenis, Blumea 28 (1982) 159.
Palynology — Pentastemona pollen is small-sized (13-23 um) and ellipsoidal to sphe-
roidal. The grains are inaperturate, but the intine shows a thick oncus, which limits the
exit of the pollen tube at germination. The exine is intectate; exine thickness is c. 0.6 urn
throughout (nexine 1/3 to 1 /2 of the sexine). The sexine consists of more or less densely
arranged composite scabrae of 0.3 to 0.5 |im in diameter, in P. sumatrana sometimes in a
fossulate pattern. These scabrae are much like the Ubisch bodies on the anther wall. Simi-
lar exine ornamentation as well as resemblance to Ubisch bodies was found in Peliosan-
thes teta (Convallariaceae).
Duyfjes — Pentastemonaceae
397
Pollen of Pentastemona is different from that of Stemonaceae in being intectate and
small (av. 16 urn versus 29, 28 and 27 um in Croomia, Stemona and Stichoneuron, re-
spectively). In terms of pollen morphology, the Pentastemonaceae and Stemonaceae do
not show any affinity.
References: Ham, R.W.J. M. van der, Blumea 36 (1991) 127-159. — Steenis, C.G.G.J. van,
Blumea 28 (1982) 159. R w ; M yan der Ham
Chromosome number — 2n = 14 (P. egregia).
Reference: Duyfjes, B.E.E., Blumea 36 (1991) 242.
Note — The descriptions of both species given below have been largely made on living
material and material preserved in alcohol.
KEY TO THE SPECIES
la. Inflorescence compound, as long as or exceeding the petiole; flowers 3-5 mm long,
tepals free 2. P. sumatrana
b. Inflorescences simple, generally shorter than the petiole; flowers 10-15 mm long,
tepals partly united into a tube 1. P. egregia
1. Pentastemona egregia (Schott) Steenis,
Blumea 28 (1982) 162; Meijer & Bogner,
Nature Malaysiana 8, 1 (1983) 26; Hotta, Rep.
1987/1988 Sumatra Research (1989), pi. 7,
lower left; Duyfjes, Blumea 36 (1991) 245, f.
2, 3. — Cryptocoryne egregia Schott in Miq.,
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 122. —
Type: Korihals s.n., West Sumatra (L, sh.
898-88.409).
Plants almost glabrous, up to 25 cm; stem
terete, 5-10 cm long, c. 1 cm diameter. Leafblade
ovate to broad-ovate, base shallowly emarginate to
cordate, apex short or distincUy acute-acuminate,
7-17 by 5-10 cm; petiole 4-6(-15) cm long. In-
florescences shordy peduncled, simple, pauciflor-
ous racemes, hidden under the leaves, 2-5 cm
long, after flowering bent towards the ground;
bracts ovate, 5-12 mm long, leaf- like, one-nerved,
the nerve smooth or with antler-like branched
emergences, margin translucent and irregularly
undulate. Flowers campanulate-urceolate, 12-20
mm long, 18-20 mm diameter; pedicels 3-4
mm long; perianth rather thickly fleshy, connate
for slightly over half of its length, pale greenish
yellow, tube 6-12 mm long, c. 10 mm wide,
lobes roundish, 6-9 mm long, spreading and
somewhat recurved in anthesis. Staminal ring
fused with the basal pan of the perianth tube and
the hypanthium: outgrowths of the connectives
narrow and somewhat bent down towards the stig-
ma; ovary dentately ribbed, shorter than the peri-
anth, c. 5 mm high, the ribs extending on the
perianth-tube; stigma flattish, 3- or 4-lobed. Fruit
c. 13 mm long, with 10 firm ribs fringed with
ander-like emergences which extend on the persist-
ing perianth. Seeds many, almost globular, c. 1.5
by 1.25 mm, glossy. - Figs. If, 2c, d.
Distribution - Malesia: Central West Sumatra,
very local.
Habitat & Ecology - On steep shady slopes in
primary forest, up to 500 m altitude.
2. Pentastemona sumatrana Steenis, Blumea
28 (1982) 161; Duyfjes, Blumea 36 (1991) 245, f.
2, 3. — Type: de Wilde & de Wilde-Duyfjes
18695, N Sumatra (L).
Herbs to 40 cm tall, with scattered hairs; stem
short, up to 5 cm. Leafblade broad-ovate, base deep-
ly cordate, apex acute, 9-16 by 6-9 cm, upper
surface glabrous, lower surface hairy on the nerves;
petiole hairy, 6-20 cm long. Inflorescence com
pound, with (1 or) 2-4 densely flowered, raceme-
like branches; bracts ovate-lanceolate, green, 3-8
mm long, one-nerved, the nerve hairy, margin shal-
lowly undulate; peduncle (sub)erect, 10-30 cm
long, hairy. Flowers ± globose in bud, opening
during anthesis and the male flowers soon falling
off: perianth 5-6 mm diameter; pedicels c. 2 mm
398
Flora Malesiana ser. I, Vol. 11 (2) (1993)
long; tepals delicate, free, broad-ovate, creamy
white, 2-3 mm long, somewhat accrescent in
fruit. Staminal ring fused with the hypanthium;
outgrowths of the connectives broad, flattish.
Ovary undulately ribbed, as long as the perianth,
c. 2.5 mm high, stigma 3- or 4-lobed, but in age-
ing flowers becoming entire, flat and round. Fruit
c. 4 mm long, with 10 ribs, with faintly undulat-
ing ridges or with antler-like emergences, not ex-
tending on the persisting perianth. Seeds up to 60,
ellipsoid, c. 1 mm long, distinctly spiny ribbed. -
Figs, la-e, 2a, b, 3.
Distribution - Malesia: N Sumatra, southern
Alas Lands; very locally.
Habitat & Ecology - Among rock blocks in
damp places, up to 100 m altitude.
Fig. 3. Pentastemona sumatrana Steenis. Rower,
c. x 10 (Photograph P.J. van der Vlugt, taken from
a cultivated specimen).
Flora Malesiana scr. I, Vol. 1 1 (2) (1993) 399-409
STEMONACEAE
(Brigitta E. E. Duyfjes, Leiden, The Netherlands)
Twining (to several metres long), trailing or erect perennial herbs, mostly with fasciculate
tubers, or with a short rhizome. Plants glabrous, or with uniseriate hairs. Leaves papyra-
ceous when dry, alternate, or opposite, or verticillate (China, Japan); blade (elliptic-)ovate
or broad-ovate, nerves basal or lateral, curved, shallowly depressed above, secondary
intervenation finely trabeculate, leaf margin entire; petiole at base pulvinate (Stemona), or
slightly sheathing (Stichoneuron). Inflorescences axillary, sessile, or peduncled cincin-
nae, appearing as short racemes, rarely one-flowered; flowers and bracts often dotted
with raphides; bracteoles absent. Flowers consisting of 4 similar segments, representing
two rows of two tepals, these free, valvate, out-curved at anthesis, persistent; pedicel
articulated. Stamens 4, epitepalous; filaments short, adnate to base of tepals, at base
mutually free or shortly connate; anthers consisting of two ovoid or elongate thecae, each
opening by a longitudinal lateral slit; the thecae situated on top of the filament, either with-
out (Stichoneuron) or with an apically enlarged tepal-like appendage of the connective,
moreover the connective with a median longitudinal ridge separating the thecae, the ridge
either smooth and thin, or fleshy and with a brain-like wrinkled structure, the thecae them-
selves in addition often protruding into a common sterile appendix, 1-8 mm long, of
which the tips may be fused, thus forming a crown-like structure over the stigma (Stemona).
Ovary superior or half superior, small, one-celled, ovules few to many, basally (Stemona)
or apically (Stichoneuron) attached, anatropous or semi-anatropous; style absent; stigma
inconspicuous, roundish, papillose. Fruit a 2-valved capsule; seeds few to many, broad-
ellipsoid, faintly or conspicuously longitudinally ridged; funicle long, with a coralloid, or
lobed, hollow, wide-celled aril (Stemona), or aril in the form of uniseriate hairs (Sticho-
neuron). Endosperm present.
Taxonomic position — An isolated family because of various special morphological
features of the flower. Its affinity has generally been accepted as being with the Liliaceae
s.l., although not closely. Burkill (1960) and Ayensu (1964) suggested an affinity with
Dioscoreaceae.
References: Ayensu, E.S., Bot. Gaz. 129 (1968) 160-165. — Burkill, I.H., J. Linn. Soc. Bot. 56
(1960) 319-412. — Dahlgren, R.M.T., H.T. Clifford & P.F. Yeo, The families of the Monocotely-
dons (1985) 121. — Duyfjes, B.E.E., Blumea 36 (1991) 239-252. — Krause, K., in A. Engler, Nat.
Pflanzenfam. ed. 2, 15a (1930) 224-227. — Prain, D., J. As. Soc. Beng. 73 (1904) 39-44. — Rogers,
G.K., J. Arnold Arbor. 63 (1982) 327-336. — Steenis, C. G.G.J, van, Blumea, 28 (1982) 151-163.
Distribution — Three genera, two of which occur in Indo-Malesia; Stemona (c. 20
species) occurs in Japan and Continental Asia, extending through Malesia to northern
Australia, Stichoneuron (2 species) in continental SE Asia and the Malay Peninsula. The
genus Croomia (c. 3 species) has a disjunct distribution in eastern China, southern Japan,
and SE United States (Florida, Georgia). The genus Pentastemona, described by Van
Steenis (1982) in Stemonaceae, has been removed from this family and raised to family-
rank (see under Pentastemonaceae). (399)
400 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Habitat & Ecology — Members of Stichoneuron and Croomia are small, delicate forest
dwellers at low altitudes. Stemona species mostly prefer a seasonal climate and can be
found in rather dry places, often rocky habitats and in seasonal forests, generally not very
far from the coast, at altitudes below 500 m.
Floral morphology — The early development of the stamens in Stemona and Sticho-
neuron is similar in spite of significant differences in the adult stages. A thorough compar-
ison of the floral morphology of Stemona, Stichoneuron and Pentastemona (see Pentaste-
monaceae, p. 395) supports the exclusion of the genus Pentastemona from Stemonaceae.
Reference: Heel, W. A. van, Blumea 36 (1992) 481-499, pi. 1-5.
Seeds — The seeds are distinctly ridged and have a well-developed raphe and chalaza.
Dispersal is probably zoochorous.
Reference: Bouman, F. & N. Devente, Blumea 36 (1992) 501-514, pi. 3-5.
Leaf anatomy — Crystals are present as styloids and raphides {Stemona), or as raphides
only {Croomia). In Stichoneuron caudatum irregular crystalline conglomerates ('clustered
needle-shaped + diamond- shaped + irregularly shaped crystals', somewhat related to
raphide bundles but not identical) are present, but styloids are absent. In Stichoneuron
membranaceum the same rather undefined crystals occur together with raphide bundles.
Reference: Baas, P., in C.G.G.J. van Steenis, Blumea 28 (1982) 159.
Chromosomes — Stemona japonica: 2n = 14 (Suzuka & Koriba 1949); Stichoneuron
caudatum: 2n = 18 and Croomia pauciflora: 2n = 24 (counted by Gitte Peterson, Copen-
hagen, unpubl.). Dahlgren et al. (1985) mentioned for the family x = 7.
References: Dahlgren, R.M.T., H.T. Clifford & P.F. Yeo, The families of the Monocotelydons
(1985) 121. — Suzuki & Koriba, Jap. J. Pharmacog. 3 (1949) 68.
Palynology — Pollen grains of Stemonaceae are small to medium-sized (19-40 u.m),
ellipsoidal to spheroidal, and monosulcate or inaperturate. The exine is distinctly stratified
(tectum, infratectal layer, nexine). Ornamentation is diverse. Exine thickness ranges from
0.5 to 1.5 u.m at the proximal side. Pollen of the Malesian species of Stemona can be easi-
ly distinguished from Stichoneuron pollen, even with light microscopy. Exine ornamenta-
tion is rugulate to scabrate in Stemona javanica, S. lucida, S. parviflora and S. tuber osa,
fossulate in S. curtisii, psilate/perforate in S. australiana, and microreticulate in Sticho-
neuron. A few Stemona species from Thailand {S. kerrii, S. phyllantha) may be difficult
to separate because of their similar microreticulate ornamentation. Stemona pollen is mono-
sulcate and Stichoneuron pollen is inaperturate. However, the latter frequently shows a
sulcus-like crack. Pollen of Stemona is also distinct on account of its granular/columellate
infratectal layer. In Stichoneuron it is columellate. Pollen of Croomia (SE United States,
Japan, China) resembles Stichoneuron pollen (reticulate ornamentation, columellate infra-
tectum, monosulcate). Pentastemona, formerly included in the Stemonaceae, is pollen-
morphologically quite different from Croomia, Stemona and Stichoneuron (see Pentaste-
monaceae, p. 396).
References: Steenis, C.G.G.J. van, Blumea 28 (1982) 151-163. — Ham, R.W.J. M. van der, Blumea
36 (1991) 127-159. R _ w> ;> M _ van der Ham
Duyfjes — Stemonaceae 401
Phytochemistry — As far as I am aware nothing is known about the chemistry of the
genera Pentastemona and Stichoneuron. The roots of several species of Stemona are used
in Southeast Asia, China and Japan as insecticides and therapeutical agents (e.g. Perry &
Metzger 1980). The chemistry and the chemotaxonomic aspects of Stemona and Croomia
were discussed twice during the past 30 years (Hegnauer 1963, 1986). In these two trea-
tises many references are available. Subterranean parts of both genera contain several in-
secticidal alkaloids which represent a special type hitherto only known from these taxa.
Tuberostemonin, C22H33NO4, is the best known of the Stemonaceae alkaloids. It was
originally isolated from roots of Stemona tuberosa and also occurs in other Stemona taxa
and is accompanied by a series of biogenetically related bases. Stems and leaves of S.ja-
ponica yielded the alkaloids stemofoline and stemospironine and roots and rhizomes of
Croomia heterosepala contain croomine. Recently roots of Stemona species growing in
the Southwest of China are investigated for insecticidal and therapeutically useful alka-
loids: Stemona mairei (Wen-Han Lin et al. 1992), S. parviflora (Wen-Han Lin et al. 1991,
1992), S. sessilifolia (Dongliang Cheng et al. 1988) and S. tuberosa (Gwangdong Prov.;
Wen-Han Lin et al. 1992).
Hitherto Stemonaceae were thoroughly investigated for alkaloids only. All other class-
es of secondary metabolites were neglected, including their phenolic compounds. Appar-
ently there is one exception to this statement. In 1974 isolation of three non-prenylated
munduserone-type rotenoids from a Thai medicinal crude drug ascribed to S. collinsae
was reported (Shiengthong et al. 1974); these compounds were called stemonacetal,
stemonal and stemonone. Later, however, Taguchi et al. (1977) investigated the same
crude drug purchased on the Bangkok market and found it to be free of alkaloids, but
yielding the known rotenoid stemonacetal and a new one, clitoriacetal. These workers
detected that the Thai medicinal crude drug used to treat skin diseases and called 'Non-
tai-yak' or 'Non-taai-yaak' has two different botanical sources, namely roots of Stemona
burkillii, collinsae and tuberosa on the one hand, and roots of Clitoria macrophylla (= C.
hanceana) on the other, pharmacognostical investigation of their crude drug sample con-
vinced them that they were working with roots of the papilionaceous substitute Clitoria
macrophylla. This makes it highly probable that also Shiengthong et al. (1974) investigated
Clitoria macrophylla and not a Stemona taxon. Therefore, statements that Stemonaceae
contain rotenoids (Shiengthong et al. 1974; Hegnauer 1986; Ponglux et al. 1987) are sus-
picious.
Saponins were recorded for roots of 5. cochinchinensis which also contain alkaloids.
Unfortunately the chemistry of these saponins is still unknown. If Stemonaceae are re-
motely connected with Dioscoreaceae (e.g. Van Steenis 1982) or belong to Asparagales
(Huber 1991) one would guess that Stemona saponins are of the steroidal type, i.e. have
a C27-sapogenin.
References: Cheng Dongliang et al., J. Nat. Prod. 51 (1988) 202. — Hegnauer, R. Chemotaxonomie
der Pflanzen 2 (1963) 150, 436-439, 495; ibid. 7 (1986) 771-773. — Huber, H., Angiospermen. Leit-
faden durch die Ordnungen und Familien der Bedecktsamer (1991), G. Fischer, Stuttgart. — Lin Wen-Han
et al., Roy. Soc. Chem. Nat. Prod. Updates No 10684 (1991); No 13445 (1992) {Stemona parviflora); No
13740 (1992) {Stemona mairei). — Lin Wen-Han et al., J. Nat. Prod. 55 (1992) 571 {Stemona tuberosa).
— Perry, L.M. & J. Metzger, Medicinal plants of East and Southeast Asia (1980), MIT Press, Cambridge,
402 Flora Malesiana ser. I, Vol. 11 (2) (1993)
Mass. (Stemonaceae, p. 397). — Ponglux, D., et al. (comp.), Medicinal plants (1987), Princess Congress
I, Bangkok (Stemona, p. 243). — Shiengthong, D., et al., Tetrahedron Letters (1974) 2015. — Steenis,
C.G.G.J. van, Blumea 28 (1982) 151. — Taguchi, H., et al., Chem. Pharm. Bull. 25 (1977) 1026.
R. Hegnauer
KEY TO THE GENERA
la. Leaves with basal nerves only. Anther dorsally attached to the base of long, petal-like
outgrowth of the connective. Ovules rather numerous, inserted at the base of the
ovary Stemona (p. 402)
b. Leaves with lateral nerves. Anther at the top of short filament, without outgrowth of
the connective. Ovules few, apically inserted Stichoneuron (p. 408)
STEMONA
Stemona Lour., Fl. Cochin. 1 (1790) 404; C.H. Wright, J. Linn. Soc. Bot. 32 (1896)
490; Prain, J. As. Soc. Beng. 73 (1904) 39; Merr., Enum. Philipp. Flow. PL 1
(1923) 202; Schltr., Notizbl. Berlin-Dahlem 9 (1924) 190; Bot. Jahrb. 59 (1925) 541;
J.J. Smith, Bull. Jard. Bot. Buitenzorg III, 6 (1924) 73; Telford, in Fl. Austral. 46
(1986) 177. — Type species: name conserved.
Twining non-woody climbers, or trailing (N Australia) or erect herbs, mostly with
perennial tubers. Leaves alternate, opposite, or verticillate (China, Japan); blades with
curved basal nerves only; petioles not sheathing. Inflorescences sessile or peduncled,
raceme-like, rarely one-flowered. Flowers: perianth small to rather large, tepals to 6 cm
long, spreading at anthesis, the inner two petals slightly wider than the outer ones. Anthers
with the thecae dorsally attached to the base of a long petaloid outgrowth of the connec-
tive, the thecae apically with or without a (conspicuous) common subulate appendix; fila-
ments proportionally short, mostly connate, forming a ring or free. Ovary superior. Fruit
pendulous, capsular, opening with two valves, few- to many-seeded. Seeds basally in-
serted, dangling on long funicles, with a coralloid or lobed, hollow aril. — Figs. 1, 2.
Distribution — Japan, Continental Asia, through Malesia to northern Australia; in Ma-
lesia: 5 species, apparendy confined to coastal areas and areas with a more or less monsoon
climate.
Note — The frequently purplish and greenish flowers have an unpleasant smell and
may attract insect pollinators.
KEY TO THE SPECIES
la. Leaves in distal parts of shoots ail opposite; basal leaves often alternate
5. S. tuberosa
b. Leaves alternate, rarely some leaves (sub)opposite 2
2a. Inflorescence peduncled 3
b. Inflorescence sessile 4
Duyfjes — Stemonaceae
403
3a. Thecae with an appendix (excluding the outgrowth of the connectivum)
4. S. lucida
b. Thecae without an appendix (excluding the outgrowth of the connectivum)
2. S. curtisii
4a. Ridge separating the thecae irregularly undulate and fleshy 1. S. australiana
b. Ridge separating the thecae smooth and thin 3. S. javanica
L Stemona australiana (Benth.) C.H. Wright,
J. Linn. Soc. Bot. 32 (1896) 4%; Telford, in
Fl. Austral. 46 (1986) 178. — Roxburghia ja-
vanica ? australiana Benth., Fl. Australiensis 7
(1878) 1. — Type: Armstrong 628, Port Es-
sington (K holo, in Hb. Hookerianum).
Stemona versteegii Schltr., Bol Jahrb. 59 (1925)
542. — Type: Versteeg 1913, Merauke (BO
holo).
Slender glabrous twiner up to 2 m. Roots tuber-
ous, dark brown, 2-10 cm long. Leaves alternate;
blade lanceolate 8-14 by 1-5.5 cm, base cuneate,
rounded, or shallowly cordate, apex acute, nerves
3-5; petiole 0.5-2.5 cm long. Inflorescence ses-
sile, 1-8-flowered, usually one or two flowers in
full bloom, bracts parchment-like, c. 2 mm long.
Flowers: pedicel 5-40 mm long; tepals blackish-
red or dark purple, 8-10 by c. 2 mm. Stamens c.
8 mm long; anthers 2-3.5 mm long, the ridge
separating the thecae fleshy, brain-like structured,
fissured and with bulging surface; appendix of
thecae absent, staminode-like, or very short, up to
2 mm long. Fruit 10-15 by 5-10 mm, with a
beak c. 3 mm, 1-6-seeded. Seeds elongate, round-
ed at the apex, 6-8 by 2-4 mm, the base sur-
rounded by an aril consisting of hollow, finger-like
appendages.
Distribution - Australia (Northern Territory and
N Queensland); in Malesia: Irian Jaya (near Merau-
ke); Papua New Guinea (S coast).
Habitat & Ecology - In savanna and savanna
forest, semideciduous bush, in Imperata fields: in
N Queensland in the margin of rain forest and in
coastal forest on sandy laterite; altitude sea level to
c. 120 m.
Notes - According to the herbarium material
seen, the inflorescences regularly produce fruits.
Superficially S. australiana resembles S. javani-
ca, but the outgrowth of the connecdvum separat-
ing the thecae in S. australiana is fleshy and of a
brain-like structure whereas in 5. javanica it is
membranous and smooth.
Brass 6039 recorded the species as rare on Daru
Island. This also holds true for Papua New Guinea
and Inan Jaya. Of the seven collections from these
areas only one is collected after World War n.
2. Stemona curtisii Hook, f., Fl. Brit. India 6
(1892) 298; Curtis BoL Mag. 48 (1892) L 7254.
— Type: Curtis 1522, Penang, waterfall (K
holo).
Stemona minor Hook, f., Fl. Brit. India 6 (1892)
298, p.p. — Lectotype: Glenie in Hb. Thwaites
3775, Ceylon, Trincomalee, see note.
Stemona tuberosa auct. non Lour.: Ridley, Mat.
Fl. Mai. Penins. 2 (1907) 86.
Glabrous twiner up to 3 m. Roots tuberous, c.
10 cm long, forming a bundle. Leaves alternate,
seldom opposite; blade ovate-oblong, 6-21 by
2.5-10 cm, base sometimes shallowly, but more
often broadly cordate, apex c. 2 cm acuminate,
nerves 11-15; petiole 4-8.5 cm long. Inflores-
cence 2-many-flowered, on a peduncle 2.5-10
cm, not fused with the petiole; bracts narrow, 6-
10 mm long. Flowers pendent; pedicels 8-10 mm
long; tepals pink, brown pink or dark brownish
red, 17-22 mm long. Stamens 13-18 mm long;
anthers 7-10 mm long, the thecae proceeding as
sterile ridges up to the top of the tepaloid out-
growth of the connectivum, and separated by a
smooth projecting ridge, 1-1.5 mm high, equally
tapering off towards the top of the tepaloid connec-
tivum. Additional appendix of the thecae absent.
Fruit 25-30 by c. 15 mm, shortly beaked, 2-6-
seeded. Seeds 18-20 by 3-3.5 mm, the acumi-
nate apex c. 5 mm, the base enveloped by an aril
consisting of hollow, finger-like appendages.
Distribution - Sri Lanka (rare), Thailand; in
Malesia: Malay Peninsula (Lepar Archipelago near
Banka; one collection seen).
Habitat & Ecology - At low altitudes, not far
from the coast, on sand and limestone; found near
the shore, on riverbanks, near waterfalls, in thickets
and scrub, also in secondary growth in rubber plan-
tations.
Notes - The flowers of S. curtisii and S. collin-
sae Craib (Thailand) are identical, but S. curtisii is
a rather tall climber while 5. collinsae is a low,
erect herb.
Ridley I.e. mentions opposite as well as alter-
nate leaves in die same plant, but the occurrence of
opposite leaves is rare; I have seen it in only one
collection. Ridley 10305.
404
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Stemona minor is based on three collections:
Wight 2821, which is S. tuberosa, the above
mentioned collection Glenie in Hb. Twaites 3775
which is S. curtisii, and Beddome 770 in BM., the
latter not seen.
3. Stemona javanica (Kunth) Engler in Engler
&Prantl, Nat Pflanzenfam. II, 5 (1878) 8;
C.H. Wright, J. Linn. Soc. Bot. (1896) 495;
J.J. Smith, Ic. Bog. 3 (1907) 107, L 244; Backer
& Bakh. f., Fl. Java 3 (1968) 154. — Rox-
burghia javanica Kunth, En. PI. 5 (1850) 288.
— Type: Zollinger Plantae Javanicae 2441 (BM
holo).
Stemona asperula J.J. Smith, Bull. Jard. BoL Bui-
tenzorg HI, 6 (1924) 76. — Syntypes: Beguin
2333, 2338 (both Halmahera).
Stemona sulensis J.J. Smith, Bull. Jard. Bot. Bui-
tenzorg III, 6 (1924) 78. — Type: Hulstijn
(Atje) 182, Mangoli Island (BO holo).
Stemona papuana Schltr., Bot. Jahrb. 59 (1925)
542. — Type: Peekel 839, New Ireland (Neu
Mecklenburg), Lemakot (B holo).
Glabrous, or slightly asperulous (Halmahera),
twiner, up to 5 m. Roots tuberous. Leaves alter-
nate; blade ovate or broadly ovate 6-18 by 2-12
cm, base cuneate to deeply cordate, nerves 7-11;
petiole 1.5-6 cm long. Inflorescence sessile or
subsessile, (1- or) few-many-flowered, bracts
parchment-like 1.5-3 mm long, imbricately ar-
ranged and finally forming a pseudo-peduncle up to
1.5 cm long. Flowers: pedicel 1-4 cm long; tepals
dark purple or purple-brown, 8-12 by 2-4 mm.
Stamens purple or dark brown, c. 8 mm long; an-
thers 2-2.5 mm long, the thecae separated by a
c. 1 mm high, wing-like ridge; appendix of thecae
0.5-1.5 mm long, fused at the tips, or appendix
staminode-like, c. 1 mm long, not fused. Fruit
15-35 by c. 10 mm, beak c. 4 mm, 3-7-seeded.
Seeds elongate, rounded at the apex, 5-16 by 2-4
mm, funicle c. 6 mm long, with an aril consisting
of many translucent, hollow, finger-like append-
ages, reaching to the base of the seed.
Distribution - Malesia: S coast of Java, Halma-
hera, Sula Islands (Mangoli, Sanana), NE Irian Jaya,
N Papua New Guinea, S coast of New Britain, and
W New Ireland.
Habitat & Ecology - Apparently occurring only
along and near the coast, up to 300 m altitude. In
Java on sandy and rocky beaches, coastal forest
(Barringtonia-formalion), scrub vegetation, river
borders and devastated forest. In the Sula Islands on
karang (limestone) and in ladangs; in Halmahera,
Irian Jaya and Papua New Guinea in rather open
primary forest, mixed lowland rain forest, forest
edges, secondary forest and grassland.
Notes - The flowers have an unpleasant smell.
In Halmahera, Weda District, extract from the
stem is given as a drink after childbirth for purifi-
cation. In Sula Sanana (Moluccas) the tubers are
used as fish-poison (Bloembergen 4372).
Specimens cultivated in the botanical garden at
Bogor (origin not indicated) have whitish tuberous
roots.
Some specimens from Halmahera have slighdy
asperulous stems and petioles.
4. Stemona lucida (R. Brown) Duyfjes, Blumea
36 (1991) 243. — Dioscorea lucida R. Brown,
Prod. (1810) 295. — Type: Banks & Solander
s.n., Endeavor R., Queensland (BM holo; left-
hand specimen).
Stemona philippensis Merr., Bull. Philipp. Gov.
Lab. 6 (1904) 16; Telford, in Fl. Austral. 46
(1986) 178. — Type: Merrill 3061, Philippines,
Masbate Island (K holo).
Slender, glabrous twiner, up to 7 m. Roots tu-
berous. Leaves alternate; blade variable in shape,
ovate, lanceolate, hastate, or linear, 4-14 by 0.5-
9 cm, with deeply cordate, truncate or narrowed
base, nerves 3-11; petiole 1-3 cm long. Inflores-
cence 3 -many-flowered; peduncle 1-9 cm long,
not fused with the petiole; bracts 2-5 mm long,
rather membranous. Flowers: pedicel 0.3-0.5 cm
long; tepals dark purple, 8-14 by 2-3.5 mm.
Stamens 8-12 mm long; anthers c. 3 mm long,
ridge separating the thecae smooth; appendix of
thecae 1.5-4 mm long, fused at the tips. Fruit
8-20 by 7-12 mm, c. 0.6 mm beaked, 2-9-
seeded. Seeds elongate, rounded at the apex, 10 by
4 mm, surrounded at the base by a vesicular aril,
c. 4 mm long.
Distribution - Australia (northern Queensland,
Cook District); in Malesia: Philippines (Masbate,
Ficao); Papua New Guinea (Normanby Island,
Milne Bay Prov., Central Prov. in the surround-
ings of Port Moresby).
Habitat & Ecology - Rather dry places in mon-
soon areas, not far inland, in scrub, open forest, dry
gallery forest, and on roadsides; at an altitude of
0-200 m.
Notes - The type material in BM exists of two
specimens, belonging to different species. The left-
hand specimen, the holotype, bears no flowers, but
the two peduncles with the characteristic persisting
pedicels leave no doubt about its identity. The
right-hand specimen, which is sterile and without
peduncles, is probably S. australiana.
Duyfjes — Stemonaceae
405
Wright, in J. Linn. Soc. Bot. 32 (1896) 495,
placed Dioscorea lucida in S. javanica. Telford
(I.e.: 180) put it aside in Stemona as a doubtfull
name, because of an sterile isotype seen by him
in NSW.
The inflorescences of the juvenile specimen in
the collection Blake 14540, Queensland (Cook Dis-
trict), are situated in the lower part of the stem, in
the axils of c. 5 mm long cataphylls. These leaves
gradually change higher up the stem into lanceo-
late and hastate leaves.
5. Stemona tuberosa Lour., Fl. Cochin. 1
(1790) 404, but cited literature p.p.; C.H.
Wright, J. Linn. Soc. Bot. 32 (1896) 494. —
Type: A specimen not found; lectotype: Rumph.,
Herb. Amb. 5 (1747) 365 "Ubi Gorita nigrum",
with illustration t. 129.
Roxburghia gloriosoides Roxb., PI. Corom. 1
(1795) 29, t. 32; Blume, En. PI. Jav. 1 (1830)
9 {'Roxburgia'), for the description, the cited
literature p.p. — Type: not known.
Roxburghia viridiflora J. Smith, ExoL Bot. 1
(1804/05) 111. — Type: Wallich 5156 (K
holo).
Roxburghia gloriosa Pers., Syn. PI. 1 (1805) 412;
Curtis Bot. Mag. 35 (1812) t. 1500; J.J. Smith,
Bull. Jard. Bot. Buitenzorg III, 6 (1924) 75. —
Type: not known.
Glabrous twiner to 4 m. Roots forming a fas-
cicle of many thick, fleshy, yellow or black tubers,
up to c. 10 cm long. Leaves opposite, but those in
proximal part of shoots often alternate; blade ovate
or broadly ovate, 9-19.5 by 3-14 cm, base shal-
lowly cordate or cordate, top acuminate, nerves 9-
13; petiole 1.5-7 cm long. Inflorescence (l-)2-6-
flowered; peduncle 2-8 cm long, free or fused with
the petiole for 5-30 mm; bracts 5-15 m long.
Flowers: pedicels 5-15 mm long; tepals outside
green or yellow-green with dark green or purple
stripes, green towards the tips, purple or brown-
red inside with wine-red stripes, 25-50 by 4-10
mm. Stamens purple, 25-40 mm long; anthers
8-15 mm long, the thecae separated by a 1-1.5
mm high and smooth ridge; appendix of thecae
5-12 mm long, tips fused. Fruit green, pendent,
40-70 by 15-20 mm, 10-20-seeded. Seeds
9-17 mm long, acumen c. 4 mm, funicle c. 8 mm
long, the base surrounded by a vesicular aril. -
Figs. 1, 2.
Distribution - Widespread; for details see under
the varieties.
KEY TO THE VARIETIES
la. Peduncle of the inflorescence axillary, free, not
fused with the petiole .... a. var. tuberosa
b. Peduncle of the inflorescence fused with the
petiole for 5-30 mm . b. var. ternatensis
a. var. tuberosa
Distribution - SE Continental Asia, Taiwan,
Hainan; in Malesia: Philippines, Flores, Ambon.
Notes - For discussion and typification of the
name 5. tuberosa, see under var. ternatensis.
Apparently the type variety is rather rare in the
Lesser Sunda Islands and Moluccas; from Flores I
have seen only four collections. The species is de-
scribed by Rumphius from Ambon, but later col-
lections are unknown.
b. var. ternatensis (J.J. Smith) Duyfjes, Blumea
36 (1991) 243. - Stemona moluccana (Blume)
C.H. Wright var. ternatensis J.J. Smith, Bull.
Jard. Bot. Buitenzorg III, 6 (1924) 73. — Type
Beguin 1682, Temate (BO holo). - Figs. 1, 2.
Roxburgia moluccana Blume, Fl. Jav. 1 (1827) 9
('Roxburghia'), for the description only. - Ste-
mona moluccana (Blume) C.H. Wright, J. Linn.
Soc. Bot. 32 (1896) 494; J.J. Smith, Ic. Bogor.
3 (1907) 111, L 245, 246. — Type: not known.
Stemona affinis J.J.Smith, Bull. Jard. Bot. Buiten-
zorg ffl, 6 (1924) 74. — Type: Beguin 1960, Hal-
mahera (BO holo).
Distribution - Malesia: Philippines, Bali (ste-
rile), Lombok, Flores, Timor, Wetar, Jamdena I.,
Morotai, Halmahera, Ceram, Bum (sterile), Am-
bon, Irian Jaya (western part).
Habitat & Ecology - At low altitudes not far
from the coast; found on loamy soils and sandy
tuff; beach vegetation, coastal forest, undergrowth
in thick scrub, primary forest along rivers, and on
cultivated ground.
Pollination - Probably by small flies, belong-
ing to the family Longhaeidae.
Taxonomy - Variety ternatensis seems related
to Stemona phyllantha Gagnep., Bull. Soc. Bot.
Fr. (1934) 147, from Thailand, but differs by the
peduncle being pardy fused with the pedole, and by
a large perianth, with the tepals 60 mm or longer.
Uses - No uses are mentioned on the field la-
bels, but Rumphius, I.e., gives extensive data on
medicinal properties.
Notes - Loureiro added to the description of
Stemona tuberosa only one reference: "Rumphius
406
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Fig. 1. Stemona tuberosa Lour. var. ternatensis (J.J. Smith) Duyfjes. a. Leaf, x 0.5; b. inflorescence, the
peduncle partly fused with the petiole, x 1; c. androecium, x 1, d. front and lateral view of stamen, x 2.5
(after Icones Bogoriensis).
Duyfjes — Stemonaceae
407
Fig. 2. Stemona tuberosa Lour. var. ternatensis (J.J. Smith) Duyfjes. a. Leafy stem, x 0.25; b. pedicel
with ovary (tepals and stamens removed), x 3; c. fruits, x 1, d. dehisced fruit, x 1 (after Icones Bogorien-
sis).
408
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Ubium Polypoides, t. 129". There has been con-
fusion in the interpretation of Rumphius' Ubium
Polypoides or Ubi Gorita. Rumphius distinguished
in Ubium Polypoides two taxa, viz.: "Album, seu
vulgare Ubium Gorita" (Ubium with the white
tuberous roots), and "Ubi Gorita nigrum" (Ubium
with the black tuberous roots). The first, "Album,
seu vulgare Ubium Gorita" is characterized i. a. by
the peduncle being partly fused with the petiole.
The habitats of this Ubium are given in detail, and
it is evident that this taxon is the same as the pres-
ent S. tuberosa var. ternatensis, which is a com-
mon plant in the Moluccas. The second, "Ubi
Gorita nigrum", clearly depicted on table 129, is
characterized i.a. by the peduncle emerging from
the leaf-axil, not fused with a part of the petiole.
Of this Ubium Rumphius noted that it is found
much less frequently than the Ubium with the
white tuberous roots. "Ubi Gorita nigrum" is the
same taxon as S. tuberosa var. tuberosa, which
was, as far as is known, never collected on Ambon
after Rumphius' time. As the type of S. tuberosa
could not be traced, I have indicated table 129, with
the appropriate description, as lectotype of S. tu-
berosa var. tuberosa.
According to Beguin 1725 from Halmahera, the
plant has a bunch of thick fleshy roots, as also de-
scribed by Rumphius, I.e. According to the label
of Museumtuin no 991 (ex Ambon) the roots are
yellow on the outside and white within.
Rumphius, I.e., wrote that in young plants the
leaves are alternate, in older plants opposite.
Rowers and open fruits have a disagreeable fetid
odour and the plant, when crushed, has an unpleas-
ant smell. Miller NGF 9712 noted skin irritation
caused by sap of the plant.
STICHONEURON
Stichoneuron Hook.f. in Benth. & Hook.f., Gen. PI. 3 (1883) 747.
Stichoneuron membranaceum Hook. f.
Type species:
Erect, low, herbs with short rhizomes, probably dioecious. Leaves alternate; blades
with curved lateral nerves; petioles slightly sheathing at base. Inflorescences raceme-like,
short, with densely set stiff pedicels. Flowers small with recurved tepals. Stamens free,
simple, without appendages, filaments longish, apically bearing an anther consisting of
two dorsifixed thecae without appendage. Ovary semi-inferior, one-celled, ovules few,
anatropous, apically attached. Fruit with one or two seeds. Seeds with a funicle, with a
flimsy aril consisting of uniseriate hairs. — Figs. 3, 4.
Distribution — There are 2 species, S. membranaceum in SE Asia, India (Assam), and
S. caudatum in Thailand and in Malesia: Malay Peninsula.
Stichoneuron caudatum Ridley, J. Str. Br.
Roy. As.. Soc. 57 (1911) 107; Steenis, Blumea
28 (1982) 151, f. 2c, e, f. — Type: Ridley 14582
(SING holo).
Slender, erect herbs, up to 60 cm, with few
branches; stem, pedicels and leaves on lower sur-
face hairy. Leaves alternate; blade elliptic, 7-12 by
2-5.5 cm, base rounded or cuneate, apex acumi-
nate; petiole 0.5-1 cm long. Inflorescence few- or
densely-flowered, with mosdy one or two flowers
in anthesis; bracts hyaline, c. 2 mm long, margins
fringed. Flowers 6-8 mm in diameter (but see
note); pedicel filiform and stiff, 3-10 mm long;
tepals dull violet or reddish purple, c. 5 mm long,
acuminate or with a filiform appendix up to 7 mm
long, the outside glabrous or sparsely hairy, the
inside glabrous, papillose, or sparsely hairy. Sta-
mens 1.5-2.5 mm long; base of filaments adnate
to tepals, glabrous, papillose or densely hairy; the-
cae without appendices. Ovary and style minute.
Fruit with thin pericarp, slightly flattened, slighUy
curved, 1 1-19 by 4-6 mm, the top caudate for a
few millimetres, green; perianth persistent. Seeds
1 or 2, broad-ellipsoid, sharply ridged lengthwise,
5-7 mm long; funicle c. 3 mm, aril consisting of
5 or 6 wide-celled appendages surrounding the seed
up to halfway. - Figs. 3, 4.
Distribution - Thailand; in Malesia: Malay Pe-
ninsula.
Habitat & Ecology - Evergreen forest; on forest
floor, always near water: in swampy forest, rocky
streams and on riverbanks; altitude up to 200 m.
Duyfjes — Stemonaceae
409
Fig. 3. Stichoneuron caudatum Ridley. Top of flow-
ering shoot with two (young) inflorescences, each
with one flower, x 2.5 (Photograph by J. Bogner).
Uses - The leaves of this species are eaten with
betel as a tonic in convalescence; the Malay name
'kayu mati hidup' or 'expectation of death'-tree,
suggests an abortifacient (Burkill, Diet. Econ. Prod.
Mai. Penins. ed. 2, 1966, 2120).
Notes - Herbarium material often lacks flowers,
whereas fruits are known from a few collections
only; the description of the fruit and seed is after
Van Steenis (I.e., who obtained the data from Dr.
R.E. Holttum), and from Bogner 1789 (culta).
The following description of the flower is based
on living material, provided by J. Bogner, Miin-
chen: Perianth c. 6 mm long, c. 12 mm in diam-
eter; tepals acute to acuminate, spreading, margins
recurved, dirty white, the outside sparsely hairy,
the inside densely set with white or carmine hairs.
Stamens c. 2 mm long, filaments white, short and
thick, densely beset with white or carmine hairs,
anthers free, yellow, without a conspicuous con-
nectivum (see also Fig. 3).
Living material testifies that the petiole bases
are slightly sheathing, a feature not easily seen in
dried material.
Henderson (SFN 29662) noted that the species
forms an almost pure stand as undergrowth in river-
bank forest along the Gua Musang R. (Kelantan).
The flowers are rather variable: tepals may be
acute or provided with a filiform appendix, the fila-
ments and inside of tepals are densely hairy, papil-
lose, or glabrous. The species is obviously closely
related to 5. membranaceum Hook, f., known only
from a few collections in eastern India (Khasia).
All flowers seem to be hermaphroditic (but it is
possible that they are functionally female or male).
Field research is needed to explain the scantiness of
well-developed ovaries and fruits in herbaria, sug-
gesting poor fertility in the natural conditions.
Fig. 4. Stichoneuron caudatum Ridley. Older in-
florescence showing the typical 'broom-like' ar-
ranged pedicels persistent after flowering; one fruit
developed; c. x 1 (Photograph by J. Bogner).
Index to scientific plant names
411
INDEX TO SCIENTIFIC PLANT NAMES
Suprageneric epithets have been entered under the family name to which they belong preceded by the indi-
cation of their rank (subfamily, tribe, etc.). Infrageneric epithets have been entered immediately under the
generic name to which they belong, preceeded by the indication of their rank (subgenus, section, etc.).
Infraspecific epithets have been entered under the specific name to which they belong preceded by the indi-
cation of their rank (subspecies, variety, forma, etc.).
Synonyms have been printed in italics. Page numbers in bold type denote main treatment; an asterisk
behind a page number denotes the presence of a figure of the concerned taxon.
Acaena L. 228, 232, 244, 297
anserinifolia (J.R. & G. For-
ster) Druce 297, 299*
elongata L. 297
Adcnilema Blume 245
fallen Blume 245, 246
Adenostoma 229, 232, 234
fasciculatum 233
sparsifolium 233
Agapanthus 376
Agrimonia L. 228, 232, 236,
244,299
ser. Eupatoriae Juz. 300
ser. Pilosae Juz. 300
blume i G. Don 299, 300
eupatoria L. 299, 300
javanita Jungh. 299
nepalensis D. Don 299,
300*
pilosa Ledeb. 236, 300
suave olens aucL 299
Alchemilla L. 230, 232, 243,
301
sect. Brevicaules 302
sect. Longicaules 302
dendroidea Zoll. & Mor. 302
indica 302
villosa Jungh. 302*
vulcanica Zoll. & Mor. 302
vulgaris L. 301
xanthochlora 236
Alliaceae 375-384
Allium L. 375, 376, 379
altaicum 378
ampeloprasum L. 377, 380,
381
anisopodium 378
ascalonicum auct. 381
totm Regel 378, 381
bakeri auct. 383
cepa L. 376, 380, 381
Cultivar group Aggrega-
tum381
Cultivar group Common
Onion 381
(Allium cepa)
var. ascalonicum Backer
381
var. typicum Backer 381
chinense G. Don 376-378,
380, 381
exsertum (Lindley) Baker
381
fistulosum L. 376, 380,
382*, 383
giganteum 377
kurrat 376
leucocephalum 378
macrostemon 376
odorum auct. 378, 383
porrumL. 376, 377, 381
x proliferum 380
x proliferum (Moench) Schrad.
ex Willd. Wakegi group
383
ramosum 378
sativum L. 377, 379, 380,
383
schoenoprasum L. 376, 380,
383
var. sibiricum 378
schoenoprasum auct. 381
senescens 378
splendens 378
stellerianum 378
tuberosum Rotder ex
Spreng. 376, 378, 380,
383
uliginosum G. Don 383
victorialis 378
vineale 377
Amaryllidaceae 353-373
subfam. Amaryllidoideae
353, 355, 357, 359
subfam. Hypoxidoideae 353,
354, 357, 366
Amaryllis 358
atamasco auct. 372
belladonna L. 371
Amelanchier 234
Amygdalus231
persica L. 349
Ancistrum anserinifolium J.R.
& G. Forster 297
Aphanes 302
Argentina 286
Aria (Pers.) Host 307, 310
Aronia312
Aruncus Silvester 233 ,
Atamasco rosea (Sprengel)
Greene 373
Brodiaea376
Buphane 355
Cerasus acuminata Wall. 329
javanica Teijsm. & Binn. 328
Cercocarpus 229, 233
intricatus 238
Chamaebatia 233
Chamaebatiaria millefolium
233
Chrysobalanaceae 228
Cliffortia 232
Clivia 358
miniata (Lindley) Bosse 371
Coleogyne 229
ramosissima 233, 235
Comarum 286
Combretum flavovirens Lauterb.
335
Cooperia 358
drummondii Herbert 371
Coriaria L. 385, 386, 387
intermedia 386-388
japonica 386-388
myrtifolia L. 385, 387-389
nepalensis 387-389
papuana Warb. 386, 388-
390*, 391
ruscifolia L. 386-388
subsp. microphylla 386
subsp. ruscifolia 386
sarmentosa 388
sinica 387
terminalis 387, 388
thymifolia 388
412
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Coriariaceae 385-391
Cotoneaster Medikus 232, 243,
307,308
angustifolius Franchet 316
depressus 234
glaucophyllus 308
lacteus W.W. Smith 240,
308
pannosa 235
vulgaris Lindley 308
Cowania mexicana 235, 238
Crataegeae 307
Crataegus glabra auct. 315
indica L. 318
serratifolia Desf. 315
Crinum L. 358, 359
amabile Dorm 359
americanum L. 354, 359
asiaticum L. 359, 360*
augustum 355
cortifolium Hallier 359
cumingii Baker 361
defixum auct. 360
fimbriatum Baker 361
giganteum auct. 360
gracile E. Meyer 359, 361
jagus (Thomps.) Dandy 361
kirkii Baker 361
latifolium L. 361
longiflorum Thunb. 361
macrantherum Engler 359
macrophyllum Hallier 359
moluccanum Roxb. 361
northianum Baker 359
rumphii Merr. 360
sanderianum Baker 361
sumatranum Roxb. 359
zeylanicum L. 361
Croomia 397, 400
heterosepala401
pauciilora 400
Cryptocoryne egregia Schott
397
Curculigo Gaertn. 358, 366
agusanensis Elmer 369
borneensis Merr. 369
brevipedunculata Elmer
369
capitulata-(Lour.) Kuntze
367, 369*
ensifolia R. Br. 370
erecta LauL 367, 369*
glabra Merr. 367
glabrescens (Ridley) Merr.
369
(Curculigo)
latifolia Dryand. 369
var. glabrescens Ridley
369
var. latifolia 367, 369*
var. megacarpa (Ridley)
Geerinck 367, 369
var. villosa (Wall, ex
Kurz) Baker 369
megacarpa Ridley 369
orchioides Gaertn. 356, 366,
367, 369*, 370
racemosa Ridley 367, 369*,
370
recurvata Dryand. 356, 367
scapigera Hallier 367
sumatrana Roxb. 369
villosa Wall, ex Kurz 369
weberi Elmer 369
Cyrtanthus 358
mackenii Hook. f. 371
pallidus Sims 371
Dalibarda calycina (D. Don)
Seringe 283
latifolia Blume 280
pyrifolia (J.E. Smith) Blume
280
Dendropoterium Svent. 301
Dichelostemma 376
Dichotomanthes 235
Digaster sumatranus Miq.
331
Dioscorea lucida R. Brown
404, 405
Docynia234
Dryas octopetala 235
Duchesnea J.E. Sm. 286, 286
chrysantha Miq. 291, 293
fragiformis J.E. Sm. 286
x hara-kurosawae Naruhashi
& Sugimoto 293
indica Focke 286, 291,
293
sundaica Miq. 296
var. hirsuta Miq. 296
Eriobotrya Lindley 243, 308
acuminatissima Nakai 318
ambigua Merr. 314
bengalensis (Roxb.) Hook. f.
306*, 307*. 309
japonica (Thunb.) Lindley
238,239,308,309
oblongifolia Merr. & Rolfe
315
philippinensis Vidal 318
Eucharis 358
amazonica Linden ex Planch.
371
Candida Planch. & Linden 371
grandiflora auct. 372
mastersii Baker 372
sanderi Baker 372
subedentata Baker 372
Eucrosia 354
Eurycles Salisb. ex Schult. 363
alba (R. Brown) F. Muell.
366
amboinensis (L.) Lindley 366
javanica Roemer 366
sylvestris Salisb. ex Schult.
366
Exochorda 229, 233
Filipendula 229, 232, 234, 236
ulmaria 236
Fragaria L. 230, 236, 244, 285
x ananassa (Duch.) Guedes
285,286
chiloensis (L.) P. Miller 288
chrysantha Zoll. & Mor. 291
indica An<k.2S6,29\
sundaica Blume 296
vescaL. 285
virginiana P. Miller 286
Galanthus 353
Germaria latifolia Presl 338
Gethyllis L. 366
acaulis Blanco 370
afra L. 366
Geum japonicum 237
montanum 236
rivale 236
Gillenia trifoliata 233
Griffinia 358
ornata Moore 372
Haemanthus pubescens auct. 360
puniceus auct. 373
Haemodoraceae 353, 357
Hagenia abyssinica 234
Hippeastrum 358
advenum Ker-Gawl. 372
ananuca 355
aulicum (Ker-Gawl.) Herbert
372
equestre (Ait.) Herbert 372
miniatum auct. 372
pratense (Poepp.) Baker 372
puniceum (Lam.) Kuntze 372
reginae (L.) Herbert 372
reticulatum (L'Her.) Herbert
372
Index to scientific plant names
413
(Hippeastrum)
rutilum Ker-Gawl. 372
splendens Herbert 372
stylosum Herbert 372
vittatum (L'Her.) Herbert 372
Holodiscus 229, 232
Hymenocallis Salisb. 354, 358,
361
adnata Herbert 362
caribaea (L.) Herbert 362
littoralis (Jacq.) Salisb. 361
x macrostephana Baker 362
narcissiilora (Jacq.) MacBr.
362
speciosa (Salisb.) Salisb.
362
tenuiflora Herbert 362
Hypoxidaceae 353, 356
Hypoxis L. 358, 370
acuminata 356
angustifolia 356
aureaLour. 370, 371*
franquevillei Miq. 370
hygrometrica Labill. 370
interjecta 356
multiceps 356
nyasica 356
obtusa 356
rooperi 356
Imperata cylindrica 355
Kageneckia 229
Kerria 229
japonica 233, 236
Lachemilla 302
Laurocerasus (Toum.) ex
Duhamel231,326
adenopoda (Koord. & Vale-
ton) Browicz 327
javanica (Teijsm. & Binn.)
Browicz 328
ndrabilis (Kalkman) Browicz
328
wallichii (Steudel) Browicz
329
Leucojum 353, 358
aestivum L. 372
capitulatum Lour. 367
Liliaceae 375
subfam. Agapanthoideae 375
subfam. Allioideae 375-377
tribe Allieae 375
tribe Brodiaeeae 375
subfam. Gilliesioideae 375
Lindleya 229, 234
Luetkea pecunata 234, 235
Lycoris 355
radiata 355
Lyonothamnus 229
Malus Miller 243, 310
domestica Borkh. 310
pumila Miller 239, 240, 310
sylvestris Miller 310
Marcetella Svent. 301
Mespilus 232
bengalensis Roxb. 309
cotoneaster L. 308
japonicus Thunb. 309
pyracanthaL. 316
Micromeles Decne. 231, 243,
307, 310
corymbifera (Miq.) Kalkman
306*, 307*, 311*
malayensis Ridley 3 1 1
Milla 376
Molineria Colla 366
capitulate (Lour.) Herbert
367
latifolia (Dryand.) Herbert ex
Kurz369
plicata Colla 369
recurvata (Dryand.) Herbert
367
sumatrana (Roxb.) Herbert
369
Narcissus 358
jonquilla L. 373
poeticus 355, 373
pseudonarcissus aucL 355,
373
tazetta L. 373
Neillia D. Don 232, 243, 245
fallax (Blume) Blume 245,
246
rubiflora 246
thyrsiflora D. Don 245,
246*
var. fallax (Blume) Kalk-
man 246
var. thyrsiflora 247
var. tunkinensis (Vidal)
Vidal245
tunkinensis Vidal 245, 247
Neviusia 229
alabamensis 233
Nothoscordum Kunth 375-377,
379,384
inodorum (Aiton) G. Nichols
384
Oemleria 233
Osmaronia 229
Padus231
buergeri 241
cerasus 235
grayana241
laurocerasus 241
padus241
polygamum 237
ssiori 241
Pancratium L. 358, 362
amboinense L. 366
biflorum 355
illyricum auct. 362
littorale Jacq. 362
longiflorum Roxb. 363
maritimum L. 355, 362, 363
maritimum auct. 362
verecundum AiL 363
zeylanicum L. 363*
Parinarium coccineum Elm. 335
Pauridia 354
Pebosanthes 395
Pentastemona Steenis 393,
399-401
egregia (Schott) Steenis 393,
394*. 395, 396*, 397
sumatrana Steenis 393, 394*,
395, 396*, 397, 398*
Pentastemonaceae 393-
398
Photinia Lindley 231, 243, 307,
312
bartlettii Merr. 3 1 1
dasythyrsa Miq. 314
davidiana (Decne.) Cardot
306*, 307*, 312, 313
integrifolia Lindley 306*,
307*, 312, 313, 314*
var. integrifolia 313
var. subdenticulata Miq.
314
var. sublanceolata Miq.
314
luzoniensis Merr. 318
notoniana Wight & Am. 313
forma grandiflora Koord.
&Valeton313
nussia (D. Don) Kalkman
312,314
prunifolia ((Hook. & Am.)
Lindley 312, 315
serratifolia (Desf.) Kalkman
312,315
serrulaia Lindley 3 1 2, 3 1 5
B prunifolia Hook. & Arn.
315
414
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Platea oblonga Korth. ex
Valeton 328
Polydontia 331
arborea Blume 329
Polylepis 232
Potaninia 229
Potentilla L. 230, 236, 244,
286
adinophylla Merr. & Perry
287,288
archboldiana Merr. & Perry
288, 289
borneensis (Stapf) Kalkman
287, 288, 294
brassii Merr. & Perry 287,
288
erecta 235, 236
foersteriana Laut. 287, 289
var. brassii (Merr. &
Perry) Kalkman 288,
289
var. foersteriana 289
var. ima Kalkman 289,
290*
var. simplex Kalkman
289
var. strigosa Kalkman
289
fruticosa 238
gorokana Kalkman 287, 290,
291
habbemana Merr. & Perry
287, 291
hooglandii Kalkman 287,
291, 292*
indica (Andr.) Wolf 228, 287,
291
indivisa Kalkman 287, 293
irianensis Kalkman 287,
293
leuconota D. Don 288
var. borneensis Stapf 288
var. papuana F. Muell.
294
leuconota auct. 288, 294
linilaciniata P. van Royen
287, 293
mangenii Kalkman 287, 294
mooniana Wight 295
var. kinabaluensis Stapf
296
novoguineensis Merr. &
Perry 295
papuana Focke 287, 288,
294
(Potentilla)
parvula Hook. f. ex Stapf
287, 294
philippinensis Merr. 295
polyphylla Wall, ex Lehm.
287, 295
var. kinabaluensis (Stapf)
Kalkman 295, 296
var. polyphylla 295, 296
reptans L. 286
simulans Merr. & Perry 287,
296
sumatrana Sojak 288
sundaica (Blume) O. Kuntze
287, 296
wilhelminensis P. van Royen
287, 296
Poteridium Spach 301
Poterium L. 301
lasiocarpum 237
Pourthiaea 307
Prinsepia 233
utilis 234
Proiphys Herbert 358, 363
alba (R. Brown) Mabberley
365*. 366
amboinensis (L.) Herbert
363, 364*, 365*, 366
Prunus L. 230, 233, 239, 243,
319
subg. Amygdalus (L.) Focke
240, 320, 349
subg. Cerasus (Miller) Focke
240, 320
subg. Laurocerasus (Tourn. ex
Duhamel)Rchder241,
320, 326
sect. Laurocerasus 327
sect. Mesopygeum 327,
329
subg. Padus (Miller) Focke
241,320,348
subg. Prunus 240, 319
A 349
acuminata (Wall.) Dietr. 329
adenopoda Koord. & Valeton
320, 323, 327, 328
amygdalus Batsch 320
arborea (Blume) Kalkman
321,324,329,330*
var. alticola Kalkman
331
var. arborea 331, 332
var. densa (King) Kalkman
321,323,331,332
(Prunus arborea)
var. montana (Hook, f.)
Kalkman 331
var. robusta (Koord. &
Valeton) Kalkman 331,
332
var. stipulacea (King)
Kalkman 331, 332
armeniacaL. 319
avium L. 235, 240, 320
B 322, 323, 350
beccarii (Ridley) Kalkman
321,324,332
brachystachya Kalkman 322,
326, 333
brassii Kalkman 322, 324,
333
C 320, 350
cerasus 240, 320
ceylanica (Wight) Miq.
326
dementis (Merr.) Kalkman
321,324,325,334
costata (Hemsley) Kalkman
322, 324, 334
D 322, 323, 350
davidiana 240
dolichobotrys (K. Schum. &
Lauterb.) Kalkman 322,
325, 326, 334, 337*
domesticaL. 234, 240, 319
forbesii Koehne 328
fragrans (Elmer) Kalkman
321,323,334,335
gazelle-peninsulae (Kaneh. &
Hatus.) Kalkman 322,
324,335
glabrifolia Kalkman 321,
326,337
glomerata (Koehne) Kalkman
347
grisea (Blume) Kalkman 325,
326, 338
var. grisea 322, 326, 338
var. microphylla Kalkman
322, 338, 339
var. tomentosa (Koord. &
Valeton) Kalkman 321,
322, 325, 326, 338,
339
jamasakura 240
javanica (Teijsm. & Binn.)
Miq. 320, 323, 328
javanica auct. 327
junghuhniana Miq. 328
Index to scientific plant names
415
(Prunus)
kinabaluensis Kalkman 322,
324, 325, 339
lamponga (Miq.) Kalkman
322, 323, 325, 340
laurocerasus L. 234, 326
laxinervis Kalkman 322, 325,
340
lusitanica 234
luzoniensis Merr. & Quis.
350
macrophylla Sieb. & Zucc.
var. adenopoda Vidal 327
mahaleb 234, 240
malayana Kalkman 321, 323,
324, 341
marsupialis Kalkman 321,
323, 324, 341, 342*
martabanica Kurz 328
var. scortechinii King 328
mirabilis Kalkman 320, 323,
328
odorata (Henderson) Whit-
more 322, 349
oligantha Kalkman 321, 323,
325, 341
oocarpa (Stapf) Kalkman
321-323,343
padus 320
papuana Koehne 328
persica (L.) Batsch 320, 322,
349
polystachya (Hook, f.) Kalk-
man 321, 323, 324, 343
pseudoadenopoda Koord 327
pulgarensis (Elmer) Kalkman
322, 325, 344
pullei (Koehne) Kalkman
322, 325, 326, 344
var. grandiflora Kalkman
344
rubiginosa (Elmer) Kalkman
321,324,334,345
salicina Lindley 319
schlechteri (Koehne) Kalkman
322, 324, 345
sclerophylla Kalkman 322,
326,346
scortechinii (King) Koehne
328
serotina 320
serrulataLindl. 240
spicata Kalkman 322, 325.
326,346
spinosa 235. 240
(Prunus)
subglabra (Merr.) Kalkman
322, 325, 347
turfosa Kalkman 321, 323,
347
tumeriana (F. M. Bailey)
Kalkman 322, 324, 325,
327,347
versteeghii Kalkman 321,
322, 324, 325, 348
wallaceana Kalkman 322,
326,348
wallichii Steudel 320, 323,
329
yedoensis 240
zippeliana Miq. 351
Purshia glandulosa 235
tridentata 235, 238
Pygeum Gaertner 230, 231,
326, 327
acuminatum 234
apoanum Elmer ex Koehne
334
apoense Elmer 334
arboreum (Blume) Blume
329,331,338
beccarii Ridley 332
blumei Teijsm. & Binn. 331
brevifolium Hook. f. 337
dementis Merr. 334
coccineum (Elmer) Elmer 335
coriifolium Ridley 340
costaium Hemsley 334
dolichobotrys K. Schum. &
Lauterb. 334
elmerianum Koehne 345
forbesii Koehne 345
fragrans Elmer 335
gazelle-peninsulae Kaneh. &
Hatus. 335
glandulosum Merr. 341
glomeratum Koehne 347
goethartianum Koehne 340
gracilipes Koehne 340
griseum Blume 338
hagenianum Gilli 334
hookerianum King 339
lampongum Miq. 340
lanceolatum Hook. f.
var. maingayi (Hook, f.)
Ridley 339
latifolium Miq. 338
var. tomentosum Koord.
& Valeton 339
latifolium (Presl) Rehder 331
(Prunus)
laurocerasus Koehne 345
macropetalum Koehne 350
maingayi Hook. f. 339
megaphyllum Elmer 335
melanocarpum Merr. & Perry
338
monticolum Merr. 344
myriandrum Merr. 343
odoratum Henderson 349
oocarpum Stapf 343
papuanum Hemsley 334
parviflorum Teijsm. & Binn.
331, 332
var. densum King 332
var. robustum Koord. &
Valeton 332
platyphyllum K. Schum. 335
polystachyum Hook. f. 343
preslii Merr. 338
var. vulgar e Koehne 338
pubescens Merr. 34 1
pulgarense Elmer 344
pullei Koehne 344
retusum Merr. & Perry 334
rigidum Koehne 334
robustum (Koord. & Valeton)
Koehne 332
rubiginosum Elmer 345
salomonense Merr. & Perry
345
schlechteri Koehne 345
scortechinii King 337
stipulaceum King 332
subglabrum Merr. 347
sumatranum (Miq.) Miq. 331
tetradenium Koehne 345
turnerianum F.M. Bailey 347
vulgare (Koehne) Merr. 338
zeylanicum Gaertn. 326
Pyracantha Roemer 243, 307,
316
angustifolia (Franch.) Schnei-
der 316
coccinea Roemer 3 16
crenulata 237
Pyrus L. 230, 234, 243, 317
sect. Micromeles Hook. f.
310
communis L. 239. 317
granulosa Bertoloni 3 1 1
mat us 310
nussia D. Don 314
pyrifolia (Burm.) Nakai 317
serotina Rchdcr 311
416
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Quillaja 229
saponaria 237
Rhaphiolepis Lindley ex Ker
243, 317
indica (L.) Lindley ex Ker
317,318
philippinensis (Vidal) Kalk-
man 306*, 307*. 317,
318
umbeUata 235, 239
Rhodotypos 229
scandens 233
Rosa L. 228, 230, 232, 239,
244,303
centifolia L. 303
cinnamomea L. 303
davurica 236, 238
laevigata 236
luciae Franch. & Rochebr. ex
Cr6pin 303, 305*
var. wichuraiana Koidz.
305
luzoniensis Merr. 305
philippinensis Merr. 305
var. depauperate Merr.
305
rugosa 236, 240
sterilis 237
transmorrisonensis Hayata
303, 305
wichuraiana Cr6pin 305
Rosaceae 227-371
subfam. Amygdaloideae 228
subfam. Maloideae 228, 307
tribus Maleae 229, 306*,
307*
tribus Pomeae 229
subfam. Prunoideae 228
tribus Pruneae 229, 319
subfam. Rosoideae 228
tribus Alchemilleae 229,
301
tribus Dryadeae 229
tribus Potentilleae 229,
232, 285
tribus Poterieae 229, 232,
297
tribus Roseae 229, 303
tribus Rubeae 229, 247
subfam. Spiraeoideae 228
tribus Gillenieae 229, 232
tribus Neillieae 229, 245
tribus Quillajeae 229
tribus Spiraeeae 229, 232,
244
Roxburghia gloriosa Pers. 405
gloriosoides Roxb. 405
javanica Kunth 404
javanica ?australiana Benth.
403
viridiflora J. Smith 405
Roxburgia moluccana Blume
405
Rubus L. 230, 236, 239, 243,
244, 247, 248
subg. Chamaebatus (Focke)
Focke 248, 283
subg. Eubatus auct. 284
subg. Idaeobatus (Focke)
Focke 248, 257, 262
subg. Malachobatus (Focke)
Focke 248, 262, 268, 285
subg. Micranthobatus
(Fritsch) Kalkman 248,
253
subg. Rubus 248, 284
sect. Eufruucosi 248
sect Chamaebatus Focke 283
sect. Idaeobatus Focke 257
sect Malachobatus Focke
268
sect. Micranthobatus Fritsch
230, 253
acuminatissimus Hassk. 253,
258
alcaefolius 268
alceaefolius 268
alceifolius Poiret 249, 250,
268, 269*
alpestris Blume 252, 258,
264
angulosus Focke 276
apoensis Elmer 266
archboldianus Merr. & Perry
252, 258
asper Wall, ex D. Don 268
asper auct. 266
boschianus Zoll. 283
banghamii Merr. 252, 259
beccarii Focke 250, 270
benguetensis Elmer 250, 25 1 ,
270
blumei Focke 272
brassii Merr. & Perry 264
calycinoides Hayata 281
calycinus D. Don 250, 283*
forma javanicus Kuntze
283
var. suffruticosus Focke
283
(Rubus)
celebicus Blume 261
chartaceus Kuntze 285
chingii 238
chrysogaeus P. van Royen
251,259
chrysophyllus Reinw. ex
Miq. 250, 271
dementis Merr. 252, 253
copelandii Merr. 252, 259
cordiformis Kalkman 257
coreanus 238
crataegifolius 238
croceacanthus L6veill6 268
cumingii Kuntze 250, 271
dendrocharis (Focke) Focke
276
diclinis F. Muell. 253, 254
var. ikilimbu P. van
Royen 256
var. novoguineensis
(Merr. & Perry) P. van
Royen 254
var. papuanus Focke 285
x dosedlae GUli 266
edanoi Merr. 285
ellipticus J.E. Sm. 238, 252,
260
elmeri Focke 281
elongatus J.E. Sm. 249, 272
var. laevicalyx Ridley 272
ferdinandi Focke 260
ferdinandi-muelleri Focke
251,260,261
foliolosus 238
fraxinifolius Poiret 251, 261
var. haightii Elmer 261
fruticosus L. 247, 276
glabriusculus Hassk. 272
glomeratus Blume 250, 272
var. gracilis King 273
glomeratus auct. 276
grewiaefolius Koord. ex
Focke 285
guttans Focke 285
hasskarlii Miq. 276
subsp. dendrocharis Focke
276
heterosepalus Merr. 250, 273
horsfieldii Miq. 264
hypargyrus Edgew. 264
idaeus 236, 239, 257
javanicus Blume 266
keleterios P. van Royen 250,
274
Index to scientific plant names
417
(Rubus)
keysseri Schltr. ex Diels 263,
264
koehneanus 237
laeteviridis P. van Roy en 260
ledermannii Focke 272, 273
var. beleensis P. van Royen
272
lineato-alpestris Nanihashi &
Sato 262
lineatus Reinw. ex Blume
251,262,264
forma pulcherrimus Focke
262
lobblanus Hook. 272
lorentzianus Pulle 252, 262
lowiiStapf252,253,263
var. panalabanensis
Naruhashi & Sato 263
lucens Focke var. dementis
(Merr.) Focke 253
luzoniensis Merr. 249, 274
macgregorii F. Muell. 252,
263
magnibracteatus Ridley 272
malvaceus Focke 250, 274
maximus Kuntze 282
mearnsii Elmer 251, 275
megacarpus P. van Royen
252, 254, 255
merrillii Focke 261
microphyllus 237
mindanaensis Focke 285
mingendensis Gilli 266
var. trichocarpa Gilli 266
moluccanus L. 249, 250,
275, 276, 277*
var. alceifolius (Poiret)
Kuntze 268
var. angulosus Kalkman
276, 277*. 278
var. austropacificus P. van
Royen 275
var. chrysophyllus (Reinw.
ex Miq.) Kuntze 271
var. dendrocharis (Focke)
P. van Royen 276
var. discolor (Bl.) Kalk-
man 275, 277*, 278
var. glabriusculus (Hassk.)
Kuntze 272
var. glomeratus (Blume)
Backer 272, 275
var. hasskarlii (Miq.)
Kuntze 276
(Rubus moluccanus)
var. lobbianus (Hook.)
Kuntze 272
var. malvaceus (Focke)
Backer 274, 275
var. maximus Kuntze 282
var. moluccanus 277*,
278
var. obtusangulus Miq.
277*, 278, 279*, 280
var. ochrascens Blume 271
var. pyrifolius (J.E. Smith)
Kuntze 280
var. ihespesiaephyllos
P. van Royen 275
moluccus latifolius Rumph
275
moluccus parvifolius Rumph.
258, 261
montis-wilhelmi P. van
Royen 251, 261, 263
moorei F. Muell. 256
moorei auct. 256
neo-ebudicus Guillaumin
251,252,264
niveusThunb. 251,264
subsp. horsfieldii (Miq.)
Focke 264
niveus Wall, ex Hook. f.
264
novoguineensis Merr. &
Perry 252, 254
papuanus Schltr. ex Diels
251,261,265
paradoxus Ridley 254
parvifolius 238
pectineUus Maxim. 250, 284
perfulvus Merr. 249, 280
philippinensis Focke 280
plicatus Weihe & Nees 284
podocarpus Kuntze 258
pulcherrimus Hook. 262
pyrifolius J.E. Smith 250,
280
robinsonii Ridley 272
rolfei Vidal 249, 280, 281
rosaefolius 265
rosifolius J.E. Smith 251,
265
subsp. sumalranus (Miq.)
Focke 266
var. coronarius Sims 266
royenii Kalkman 252, 256
var. hispidus Kalkman
256
(Rubus royenii)
var. ikilimbu (P. van
Royen) Kalkman 256
var. royenii 256, 257
rugosus J.E. Smith 275
satotakashii Naruhashi &
Cheksum 262
schefferi Focke 245, 246
smithii Backer 250, 282
sorsogonensis Elmer 250,
282
suavissimus 238
sumatranus Miq. 251, 266,
267*
sundaicus Blume 250, 282
var. discolor Blume 275
sundaicus auct. 272, 282
tagallus Cham. & Schlech-
tend. 266
trifidus 237
trigonus Kalkman 253, 257,
285
tsiri P. van Royen 254
tsiri auct. 256
vanoverberghii Merr. 273
var. pileanus Focke 273
wichurae Focke 275
woitapensis P. van Royen
260
Sanguisorba L. 230, 232, 236,
244,301
minor Scop. 236-238, 301
officinalis L. 235-237, 301
Sarcopoterium spinosum 238
Scadoxus 358
multiflorus (Martyn) Raf.
373
Sieversia 232
Sorbaria 233, 234
Sorbeae 307
Sorbus L. 239
sect. Aria 234
sect. Aucuparia 234, 240
secL Micromeles Rehdcr
310
aucuparia 234, 239
corymbifera (Miq.) Hiep &
Yakovlev311
Spiraea L. 230, 243, 244
cantoniensis Lour. 237, 239,
244
japonica 238
koreana 238
prunifolia 233
salicifolia L. 244
418
Flora Malesiana ser. I, Vol. 11 (2) (1993)
Sprekelia 358
formosissima (L.) Herbert
373
Stemona Lour. 397, 399-401,
402
affinis J.J. Smith 405
asperula J.J. Smith 404
australiana (Benth.)
C.H.Wright 400, 403
burkillii 401
collinsaeCraib401,403
curtisii Hook. f. 400, 403
japonica 400, 401
javanica (Kunth) Engler 400,
403, 404, 405
kerrii 400
lucida (R. Brown) Duyfjes
400,403,404
mairei401
membranaceum Hook. f.
409
minor Hook. f. 403, 404
moluccana (Blume)
C.H. Wright 405
var. ternatensis J.J. Smith
405
papuana Schltr. 404
(Stemona)
parviflora 400, 401
philippensis Merr. 404
phyllantha Gagnep. 400, 405
sessilifolia 401
sulensis J.J. Smith 404
tuberosa Lour. 400-402,
405
var. ternatensis (J.J.Smith)
Duyfjes 405, 406*,
407*
var. tuberosa 405
tuberosa auct. 403
versteegii Schltr. 403
Stemonaceae 399-409
Stenomeris 395
Stenomesson elwesii 353, 354
Stichoneuron Hook. f. 397,
399-402,408
caudatum Ridley 400, 408,
409*
membranaceum Hook. f. 400,
408
Stranvaesia Lindley 307, 312
ambigua Nakai 314
davidiana Decne. 313
glaucescens Lindley 312
(Stranvaesia)
integrifolia Stapf 3 1 3
nussia (D. Don) Decne. 314
Trichopus 395
Tristagma uniflora 378
Triteleia 376
Vaccinium corymbiferum Miq.
311
Vallota 358
speciosa (L. f.) Dur. & Schinz
373
Vauquelinia 229
Woronowia 232
Zephyranthes 358
atamasco (L.) Herbert 373
brachyandra (Baker) Backer
373
Candida (Lindley) Herbert 373
carinata Herbert 373
citrina Baker 373
grandiflora Lindley 373
mesochloa auct. 373
rosea (Sprengel) Lindley 373
tubispatha (L'Her.) Herbert
373
verecunda Herbert 373
Zygalchemilla 302
3 5185 00202 7041
Index to revised families in Series I (Spermatophyta)
Aceraceae 4: 3, 592
Actinidiaceae s.s. 4: 37
Aizoaceac 4: 267
Alismataceae 5: 317; 6: 915
Alliaceae 11: 375
Alseuosmiaceac 10: 335
Amaranthaceae
4: 69. 593; 6: 915; 8: 549
Amaryllidaceae 11: 353
Anacardiaceae 8: 395
Ancistrocladaceae 4 : 8
Aponogetonaceae 4: 11; 7: 213
Araliaceae-I 9: 1, 553
Araucariaceae 10: 419
Aristolochiaccae 10: 53
Balanophoraceae 7: 783; 9: 554
Basellaceae 5: 300
Bat(id)aceae S: 414
Betulaceae S: 207; 6: 917
Bignoniaceae 8: 114; 9: 554
Bixaceae s.s. 4: 239
Burmanniaceae
4: 13, 592; 5: 553; 9: 554
Burseraceae 5: 209. 567
6: 917; 7: 820; 9: 555
Butomaceae 5: 118
Byblidaceae 7: 135
Callitrichaceae 4: 251
Campanulaceae
6: 107. 928; 9: 556
Cannab(iii)aceae 4: 222
Cappar(id)aceae 6: 61; 7: 822
Caprifoliaceae
4: 175; 6: 928; 9: 556
Cardiopieridaccae 7: 93
Celastraceae 6: 227. 389. 930
Cenlrolepidaceae S: 421
Ceratophyllaceae 4: 41
Chenopodi aceae
4: 99. 594; 6: 932; 9: 557
Chloranthaceae 10: 123
Chrysobalanaceae 10: 635
Clethraceae 7: 139
Cochlospermaceae 4: 61
Combretaceae
4: 533; 5: 564; 6: 932
Conifcrales 10: 337
Connaraceac
5: 495; 6: 933; 9: 557
Convolvulaccac 4: 388. 599
5: 558; 6: 936; 7: 823; 9: 558
Coriariaccae 11: 385
Comaceae 8: 85
Corynocarpaceae 4: 262; 5: 557
Crassulaceae 4: 197; 9: 558
Crucifcrac 10: 541
Cryplcroniaccac 8: 187
Ctcnolophonaceae 10: 629
Cupressaceae 10: 442
Cypcraccae 7: 435; 9: 107
Datiscaceac 4: 382
Dichapetalaceae 5: 305; 6: 941
Dilleniaceae 4: 141; 7: 824
Dioscoreaceae 4: 293
Dipsacaceae 4: 290
Diplerocarpaceae 9: 237
Droscraceae
4: 377; 5: 557; 9: 562
Elaeagnaccae 10: 151
Elalinaceae 4: 203
Epacridaceae 6: 422
Ericaceae 6: 469, 943
8: 549; 9: 562; 10: 716
Erythroxylaceae 5: 543; 8: 549
Fagaceae 7: 265; 9: 563
Flacourtiaceae 5: 1, 565
6: 943; 7: 827; 9: 563
Flagellariaceae 4: 245; 9: 564
Geraniaceae 6: 445; 9: 565
Gnetaceae 4: 336; 6: 944
Gonystylaceae 4: 349
Goodeniaceae 5: 335, 567
6: 949; 7: 827; 9: 566
Haemodoraceae 5: 111; 10: 717
Haloragaceae 7: 239, 828
Hamamelidaceae 5: 363
Hippocraleaceae 6: 389
Hydrocharitaceae 5: 381
6: 952; 7: 828; 9: 566; 10: 717
Hydrophyllaceae 4: 207
Hypericaceae 8: 1; 10: 717
Icacinaceae 7: 1; 9: 566
Iridaceae 8: 77; 10: 717
Ixonanthaceae 10: 621
Juglandaceae 6: 143
Juncaceae 4: 210; 9: 566
Juncaginaceae 4: 57
Labiatae 8: 301; 9: 566
Legum.-Mimosoideae 11: 1
Leeaceae 7: 755
Lemnaceae 7: 219
Lentibulariaceae 8: 275
Liliaceae s.s. 9: 189
Linaceae 10: 607
Loganiaceae 6: 293. 953; 9: 567
Lophopyxidaceae 7: 89
Magnoliaceae 10: 561
Malpighiaceae 5: 125
Martyniaceae 4: 216
Menispermaceae 10: 157
Mimosaceae 11: 1
Monimiaceae 10: 255
Moringaceae 4: 45
Myoporaceae 4: 265
Myricaceae 4: 276
Najadaceae 6: 157
Nyctaginaceae 6: 450
Nyssaceae 4: 29
Ochnaceae 7: 97
Olacaceae 10: 1.717
OnagTaceae 8: 98
Opiliaceae 10: 31
Oxalidaceae
Papaveraceae
Passifloraceae
Pedali aceae 4
Pentaphragmataceae
Pentaphylacaceae
Pen tastemon aceae
Philydr aceae
Phytolaccaceae
Pinaceae
Pittosporaceae 5
Plumbaginaceae
Podocarpaceae
Pod os icm aceae
Polemoni aceae
Polygal aceae
Pontederiaceae
Portulac aceae
Primulaceae
Proteaceae
Punicaceae
Restionaceae
Rhizophoraceae
5: 429; 6:
R os aceae
Sabiaceae
Salic aceae
Salvadoraceae
S arcosperm a( ta)ceae
Saurur aceae
Scyphostegiaceae 5:
Simaroubaceae
Sonnerati aceae
4
Spargani aceae
Sphenocleaceae
Sphenostemonaceae
Stackhousiaceae
Staphyleaceae
Stemonaceae
Stylidiaceae 4
Styracaceae 4
Symploc aceae
8: 205; 9:
Taccaceae
Tax aceae
Thymelaeaceae
4: 349;
Trapaceae
Trigoni aceae
Trimeni aceae
Triuridaceae
Tumeraceae
Typhaceae
Ulm aceae
Umbelliferae
5: 555; 6: 983; 7:
Valcrianaceae
Violaceae 7: 179.
Xyridaceae 4: 366
Zygophyllaceae
7: 151,829
5: 114
7
216; 7
4
5
405
829
517
121
280
4:
6: 1
4:
11: 393
4: 5
4: 228
10: 447
345; 6: 960
4: 107
10: 351
: 65; 6: 963
4: 195
10: 455
4: 255
7: 121
6: 173
5: 147
4: 226
5: 416
965; 8: 550
11: 227
10: 351
5: 107
4: 224
4: 32
4: 47
297; 6: 967
6: 193. 968
513; 6: 973
233; 10: 718
4: 27
10: 145
4: 35
6: 49
11: 399
529; 6: 976
: 49; 9: 568
569; 10: 718
7: 806
10: 347
976; 7: 830
4: 43
4: 59
10: 327
10: 109
4: 235
242; 6: 982
8: 31
4: 113.595
830; 9: 569
4: 253
831; 10: 720
598; 9: 571
4: 64
Cyalhcaceae
Gleicheni aceae
lsoctaceae
Index to revised families in Series II (Pteridophyta)
1: 65
Lindsaea group
1: 177
Tectaria group
1: 1
Lomariopsis group
1: 255
Thelypteridaceae
1: 62
Schizaeaceae
1: 37
2: 1
1: 331
ISBN 90-71236-19-6
FLORA MALESIANA
Compiled and published under the auspices of
Foundation Flora Malesiana
Editorial Committee:
C. Kalkman P.F. Stevens
D.W. Kirkup W. J. J.O. de Wilde
H.P. Nooteboom
Board of the Foundation:
Aprilani Soegiarto, Jakarta, chairman
P. Baas, Leiden, vice-chairman
M.C. Roos, Leiden, secretary / treasurer
B. A. Barlow, Canberra
K. Iwatsuki, Tokyo
K. Larsen, Aarhus
A. Latiff Mohamad, Bangi
G.LI. Lucas, Kew
D. A. Madulid, Manila
Ph. Morat, Paris
F.S.P. Ng, Rome
D.H. Nicolson, Washington
P. H. Raven, St. Louis
M. A. Rifai, Bogor
S.H. Sohmer, Forth Worth
P.F. Stevens, Cambridge, USA
Requests for scientific information to be addressed to the Editors,
at the address below.
Subscription orders and requests for sample copies to be sent to:
Publication Department
Rijksherbarium / Hortus Botanicus
P.O.Box 9514
2300 RA Leiden
The Netherlands Fax (3 1) (7 1) 2735 1 1
Saturday, January 9, 2016
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