THE GARDENS’ BULLETIN, SINGAPORE

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conservation, with emphasis on the plant life of the Southeast Asian-Pacific region.

Dr. K.M. Wong Dr. Jana Leong-Skorniékova

Singapore Botanic Gardens Singapore Botanic Gardens

(Editor) (Assistant Editor)

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National Parks Board, Singapore Singapore Botanic Gardens

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Singapore Botanic Gardens National Parks Board

National Parks Board, Singapore Singapore

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Yale University Singapore Botanic Gardens

U.S.A. National Parks Board, Singapore

Dr. Rogier P.J. de Kok Dr. J.F. Veldkamp

Royal Botanic Gardens, Kew, Nationaal Herbarium Nederland

U.K. The Netherlands

Dr. W.J. Kress Dr. Jun Wen

National Museum of Natural History National Museum of Natural History

Smithsonian Institution, U.S.A. Smithsonian Institution, U.S.A.

Dr. David J. Middleton Professor Nianhe Xia

Royal Botanic Garden Edinburgh South China Institute of Botany

U.K. PR. China

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(continued on inside back cover)

IV » AUG O1 201 ARNOLD ARBORETUM

~ LIBRARY

The Gardens’ Bulletin

Singapore

VOL. 62 (2) 2011 ISSN 0374-7859

CONTENTS

R. Camara-Leret and J.F. Veldkamp

A remarkable new Medinilla (Melastomataceae) from Celebes (Sulawesi), Indonesia .........213

W.L. Goh, S. Chandran, K. Kamiya and K.M. Wong

A natural hybrid between Dendrocalamus pendulus and Gigantochloa scortechinii

(Poaceae: Bambusoideae: Bambuseae) in Peninsular Malaysia ...............:::ccssseeeseeeeeeeseeenee 229

Mark Hughes and Deden Girmansyah

A revision of Begonia sect. Sphenanthera (Hassk.) Warb. (Begoniaceae) from Sumatra ...... 239

R. Kiew and C.L. Lim

Names and new combinations for Peninsular Malaysian species of Codonoboea Ridl.

OUR ESRD) oS SS Ea UG ee na eo nL ee ee ee nee

Kiaw Kiaw Ng, P.C. Boyce and S. Othman

Studies on Homalomeneae (Araceae) of Peninsular Malaysia I: An historical and taxonomic

review of the genus Homalomena (excluding Chamaecladon) ..........ccccccecceccescesceseeseeseeeens 277

V.D. Nguyen and P.C. Boyce

Pycnospatha (Araceae: Lasioideae), a new generic record for the flora of Cambodia and

ARUP EIEN oc occcecdbcet Stan toSe See ee RR EE CEE ae aoe a 291

C. Rattamanee, S. Sungkaew and Y. Paisooksantivatana

Additional notes on Maclurochloa montana (Poaceae: Bambusoideae) in Thailand ............ 297

J.F. Veldkamp

The nomenclature of Uvaria velutina Roxb. ex Blume (Annonaceae) ................:::cseeseeeees 301

K.M. Wong

Steenisia (Rubiaceae) newly recorded for mainland Asia with a new variety of

BME NEAACT TIGR (NUTS HIAW)) ES ARGR Le cn con cosa coe wnnnnnennnmecnce ent ocecteccescreceeesnnaencnranaceddectesenacesenrstsnss 307

Sin Yeng Wong, P.C. Boyce and B.A. Fasihuddin

Studies on Homalomeneae (Araceae) of Borneo III: The helophytic Homalomena of Sunda ... 313

T.L. Yao

Ginalloa siamica vat. scortechinii is a species of Viscum (VisCaceae) ..........:::cceeeeeeeseereere 327

BOOK REVIEWS

Trees of Tropical Asia. An Illustrated Guide to Diversity James V. LaFrankie, Jr. 2010. (George

Staples) .ccccccslecasccessccusesectenceasebeeoanecct cescUeaOhe cet teete date tte tes ahaa enantio an 329

Flora of Peninsular Malaysia. Series II: Seed Plants, Volume 1. R. Kiew, R.C.K. Chung, L.G.

Saw, E. Soepadmo & P.C. Boyce (eds). 2010. (Paul K.F. Leong & K.M. Wong) ............... 331

Malaysia Plant Red List, Peninsular Malaysian Dipterocarpaceae. L.S.L. Chua, M. Suhaida, M.

HamidahGalnGasawae OlON (atoll erty euizayiel)) peeeeeeeeeeee eset 333

Alexander von Humboldt and the Botanical Exploration of the Americas. H. Walter Lack. 2009.

(George Staples): ..c...csccccsecssscensonaeseoves deusanassooast se cacssoucblascd oe tetera eae aurea ea aan eee 335

Bamboos at TBGRIVK-C. Koshy, 2010 (KIMI W O10 G))eresrere reenter 337

Date of publication: March 2011

Published and copyrighted by

National Parks Board

Singapore Botanic Gardens

1 Cluny Road

Singapore 259569

Printed by Oxford Graphic Printers Pte Ltd

Gardens’ Bulletin Singapore 62 (2): 213-221. 2011 213

A remarkable new Medinilla (Melastomataceae)

from Celebes (Sulawesi), Indonesia

R. Camara-Leret' and J.F. Veldkamp”

Netherlands Centre of Biodiversity — Naturalis,

section National Herbarium of The Netherlands, Leiden University

PO Box 9514, 2300 RA Leiden, The Netherlands

'rcamaraleret@gmail.com

*veldkamp@nhn.leidenuniv.nl (corresponding author)

ABSTRACT. A new species of Medinilla (Melastomataceae) from Celebes, Indonesia, is

described. It is a terrestrial shrub or treelet with specialised leafless inflorescence-bearing,

many-branched branches at the base of the boles. It is clearly distinct in habit, branch-, leaf-,

flower- and fruit morphology from the few other species that are reported to have this type of

flowering.

Keywords. Celebes, Medinilla, Melastomataceae

Introduction

Dr. J.B.K Arsana (then of the Kebun Raya Eka Karya, Bali) in July 1996 found a

curious species of Medinilla Gaud. (Melastomataceae) near the village of Dani in the

Bukit Batu Setan Forest, near Enrekang, in the northern part of the S Celebes Province,

Indonesia, at 550 m altitude (E/9960709, EK, L). Two cuttings were brought to the

Kebun Raya Eka Karya (“EK”) (planted at plot no. XV.B—242), at c. 1250 m alt.

and from these all subsequent collections have been made. JF V and Dr. M.M.J. Van

Balgooy (L) were kindly allowed to make a few cuts of branchlets, inflorescences,

flowers, and fruits [Van Balgooy 7520 (spirit), 7557, Veldkamp 8943, also in spirit;

L; collections from one of the two are J Made Suja HK. 826 in EK, June 19th, 2007].

They are terrestrial erect shrubs with inflorescences borne on specialised

leafless, incrassate, many-branched branches at and above ground level and sometimes

somewhat higher up on the stems forming a dense mat around the base of the plants

(Fig. 1).

Mr. A. Kurniawan (EK) and Dr. M.M.J. van Balgooy observed that they flower

and fruit throughout the year. Apparently well-developed seeds are produced in great

numbers, but although attempts were made, none have germinated. Cuttings, however,

were quite successful.

Inflorescence terminology

It is not quite clear what this kind of inflorescence is to be called. The terminology for

inflorescences is quite confusing and sometimes contradictory. The type encountered

here is a special kind of cauliflory (Fig. 2). Various terms have been suggested.

Geocarpy is too broad a term as it refers to all kinds of situations where diaspores are

214 Gard. Bull. Singapore 62 (2) 2011

associated with soil. Rhizocarpy is erroneous, as the flowers are nor from specialised

roots.

Flagelliflory is ambiguous as it is used in two meanings. Weberling (1989:

233-234) defined flagelliflory as when “the flowers or inflorescences hang down from

the crown of the tree on long rope-like branches, and so are brought into an exposed

position which allows the flowers (and fruits) to be visited by bats”, e.g., in the African

Sausage tree, Kigelia africana (Lam.) Benth., Bignoniaceae.

In another usage, especially favoured by students of the Annonaceae and

Moraceae, the term refers to specialised whip-like leafless branches arising more-or-

less at ground level and spreading away, often covered by soil and fallen leaves for

sometimes more than 10 m from the stem (Mildbread 1922: 116, Schatz & Wendt

2004). The latter approaches the present situation, were it not that the branches are

specialised short shoots (brachyblasts), and not whip-like but more sausage-like, and

do not spread so far. Van der Pijl (1982: 26—28) called it basicaulicarpy (fruits at the

base of the bole), but this does not exclusively indicate the presence of the brachyblasts.

Perhaps the nearest in meaning is what Mildbread (1922: 116) has called

idiocladanthy (flowers on special branches): plants that flower on special leafless

branches with distinct internodes, where related species have axillary solitary flowers.

Curiously, Mildbread states that species with axillary panicles “of course” do not

belong here, but does not explain what in his eyes the fundamental difference between

solitary flowers and panicles would be.

Fig. 1. Medinilla tapete-magicum Camara-Leret & Veldk. in the Kebun Raya Eka Karya, 15

January 2010. Note the other specimen in the back behind JFV. Photo credit: Mr. A. Kurniawan.

Anew Medinilla from Celebes DUS

Methods

The term “torus” in the flowers as used here is in the sense of Gleason (1939) for the

ring of the hypanthium above the ovary upon which the petals and stamens are inserted.

The length of the hypanthium was measured from the apex of the torus downwards to

the attachment to the pedicel. Its width was taken at the middle of its length. Sepals

were measured from the torus upwards. The width of the buds was taken at the widest

part of the petals.

Discussion

Medinilla is the largest genus of Melastomataceae in Malesia (c. 360 spp., with an

endemicity of nearly 92%: Bodegom & Veldkamp 2001). The latest revision of the

species of the family in the Dutch East Indies is by Bakhuizen van den Brink f. (1943,

1946, 1947), in which this species could not be found. More-or-less recent local

revisions are available for Thailand (Renner et al., 2001), Java (Backer & Bakhuizen

van den Brink f. 1964), while those for Borneo and The Philippines were treated by

Regalado (1990, 1995). For the last area, he listed four species apparently with a

similar flowering habit.

Medinilla aurantifiora Elmer (1911) from Negros, Panay, and Sibuyan is also

an erect terrestrial shrub, as documented by Regalado (1995: 153, t. 11). It differs

by the branchlets having soft, yellow, stellate to plumose deciduous hairs, auriculate

leaf bases, axillary inflorescences clustered on woody tubercles, 4-merous flowers,

hypanthia that are 6-7 mm long, pink to orange red petals, subequal stamens with

larger filaments, and fruits that are crowned by persistent calyx lobes. Elmer described

the inflorescences as “from near the base of the stem or stems, clustered on gnarly

ligneous tubercles 1.5 dm long, more or less branched, the ultimate one ascendingly

curved” and compared it to M. cauliflora Merr. (1910; non Hemsl., 1895). Merrill

described the inflorescences as “on the trunk below the leaves, of very short, congested

branches, forming a compact mass c. 8 cm diameter or less, the branches stout, each

bearing many, subdistichous, crowded, oblong-ovate, 9-10 mm long bracts which

are acute or obtuse, some empty, some subtending flowers”. Altogether this is quite

different from our species. Because of the homonymy Merrill’s species was renamed

M. trunciflora Merr. (1913) and synonymised by Regalado (1995: 153, t. 11!) with M

aurantiflora.

Medinilla polillensis C.B.Rob. (Robinson 1911a) from Bohol, Catanduanes,

Leyte, S Luzon, and Polillo, is terrestrial, but scandent, rooting in the nodes,

branchlets quadrangular, winged, leaves with the transverse veins adaxially faintly

distinct, abaxially absent, inflorescences cauline, the flowers more often 3, disposed

umbellately, or in two whorls each of three, flowers 4-merous, hypanthia 7-8 mm

long, petals c. 12 mm long, filaments c. 6 mm long, anthers c. 6 mm long.

We were not able to study M. pinnatinervia Merr. (Merrill 1913) from Luzon

(Cagayan, Pamplona) [lectotype: Ramos 7483, designated by Regalado (1995: 152),

isotype in US: 00344975]. It obviously is quite different, too, by being an epiphytic

scandent shrub, leaves anisophyllous, 9(—11)-pli-nerved, the larger ones distinctly

petiolate, 20-25 =x 8-10 cm, the smaller ones subsessile, c. 10 x 4 cm, and fruits

crowned by persistent calyx lobes. Merrill described the inflorescences as “caulifiorous,

216 Gard. Bull. Singapore 62 (2) 201]

racemose” and “from the stem below the leaves, of many-flowered, short, rather

dense, solitary or fascicled racemes 5 cm long or less”. He compared it to M. loheri

Merr. (Merrill 1913) and M. disparifolia C.B.Rob. (Robinson 1911b). Unfortunately

its flowers are unknown, and because of the pinninerved leaves it is possibly not a

Medinilla at all. Regalado (1995: 152) noted that he had seen four collections, also

all without flowers, and accepted it without a query as belonging to Medinilla. He

described the inflorescence as “fascicled on old wood”.

Medinilla radiciflora Quisumb. & Merr. (Quisumbing & Merrill 1928) differs

by the young branches being densely stellate tomentose, and having 11—14-pli-

nerved leaves that are 15—24 = 8.5—13.5 cm in size, with an obtuse to cordate base.

Its inflorescences are cymose, apparently borne on the roots (?), erect, densely and

many-flowered, repeatedly dichotomously branched, with peduncles up to 7 cm long.

The flowers are 4-merous, the buds c. 16 mm long, the hypanthium 8—10 mm long,

with marginal teeth 1.25—1.5 mm long, petals 10-11 x 8 mm, purplish, and stamens

of unequal length.

The new species here is also not to be confused with M. radiciflora C.Y.Wu

(1979) from Yunnan. The inflorescences of that species, judging from the illustration,

are reminiscent of our Medinilla, but it is Pseudodissochaeta lanceata M.P.Nayar from

S. China (Hainan, Yunnan). The record for Vietnam by H6 (1992; “‘/anceolata’) might

well be an undescribed species.

Medinilla tapete-magicum Camara-Leret & Veldk., sp. nov.

Frutex vel arbuscula terrestris erectus. Ramuli subtereti. Foliorum axillae setis

furfuraceis, plantae cetera glabrae. Folia opposita isomorpha longe petiolata, laminis

ellipticis ad oblongis 7-plinervis, basi acuta. Inflorescentiae densae ramis specialibus

efoliatis incrassatis multibrachiatis e caulis basi vel supra ortae interdum aliquantum

superiores plus minusve tegetes circumcirca formantes. Flores 5-meri. Hypanthium

urceolatum dentibus marginalibus minutis ad absentibus. Fructus tenuiter vallati

subglobosi 6-7 mm diam. TYPE: Van Balgooy 7557 (holo L), Eka Karya Botanical

Gardens, Bali (plot no. XV.B—242), 7 May 2007.

Erect terrestrial shrub or treelet, c. 1.5 m tall, twigs subterete, lenticellate, younger

branches ferruginous, old branches (when dry) white and lenticellate, leaf axils with

furfuraceous bristles, plant otherwise glabrous. Petioles terete, 2.5—5 cm x 2 mm,

lenticellate. Leaves opposite, subequal, elliptic to oblong, 8-17 <x 4—8 cm, 3—6 times

as long as the petioles, base acute, margin entire, glabrous, apex acute to acuminate,

texture pergamentaceous, above glaucous, underneath laxly ferruginous along veins

and reticulations, 7-plinerved, distalmost primary nerves 1.5—3 mm away from the

preceding pair, submarginal vein conspicuous, secondary nerves conspicuous on both

surfaces, angle of divergence from midvein right angle or nearly so, tertiary nerves

conspicuous. Inflorescences borne on specialised leafless incrassate many-branched

branches up to 50 cm long at and above ground level and sometimes somewhat

higher up on the stems forming a kind of dense mat of more than | meter diameter

around the base of the plants. Bracts not found. Bracteoles minute. Flowers borne

in dense basiflorous inflorescences of many-flowered cymes. Peduncles 15-20 mm

long. Pedicels in fruit 7-10 mm long. Mature floral buds 6-7 < 2-3 mm. Flowers

Anew Medinilla from Celebes Zit

5-merous. Hypanthium urceolate, 2 x 2 mm, covered with crystals (cystoliths). Torus

1-1.5 mm long, marginal teeth minute or absent, apex glabrous. Petals obliquely

diamond-shaped, 4-5 x 2 mm, pink, glabrous, with conspicuous veins. Stamens 10,

all of equal length. Filaments 2 mm long. Anthers 3 mm long, opening with | terminal

pore, lateral appendages recurved in anthesis, 0.5—1 mm long, dorsal spur rectangular,

0.75—1 mm long. Style 4 mm long, glabrous, stigma punctate, glabrous. Fruit globose,

6—7 mm diam., purple, fleshy, glabrous, inside filled with jelly; calyx teeth absent.

Seeds numerous, c. 0.5 mm long, surface minutely punctate. (Fig. 1—5)

Distribution. Sulawesi, Enrekang, Bukit Batusetan, c. 550 m alt. (c. 3°33’S 119°46’E),

cultivated in the Eka Karya Botanical Garden, Bali, plot XV.B—242, c. 1250 m alt.

Habitat. Rain forest, on clayey soil, 550(—1250) m alt.

Collectors ‘notes. Shrub in February 2004 c. 1 m tall (Van Balgooy 7520, L), in January

2010 c. 1.5 m (Veldkamp 8943, L), innovations brown pubescent, twigs lenticellate,

inflorescences basiflorous, flowers reddish white, pink, anthers yellow, fruits red

turning purple, fleshy, juicy.

Eponymy. The epithet tapete-magicum is for the “magic tapestry” formed by the

inflorescences at the base of the stems.

Fig. 2. Medinilla tapete-magicum Camara-Leret & Veldk. Close-up of inflorescences. Photo

credit: Irawati.

218 Gard. Bull. Singapore 62 (2) 2011

Fig. 3. Medinilla tapete-magicum Camara-Leret & Veldk. Branchlets and leaves. Photo credit:

J.F. Veldkamp.

Fig. 4. Medinilla tapete-magicum Camara-Leret & Veldk. Flower, bud, and very young fruits.

Photo credit: J.F. Veldkamp.

219

3. Medinilla tapete-magicum Camara-Leret & Veldk. A.

-pams alone a branch.

inflorescence. C. Flower bud. D. Open flower E. Flower, 2? petals and 4 stamens

Leaf-

Temoved. A_C—E- E19960709 (L). B: Veldkamp 8943 (L). Drawn by- A. Walsmit Sachs-Jamsen.

B. Partial

i=)

Gard. Bull. Singapore 62 (2) 2011

ACKNOWLEDGEMENTS. Dr. M.M.J. van Balgooy (L) is thanked for providing material,

advice, assistance, and lodging. Mr. A. Kurniawan and the other staff members of the Kebun

Raya Eka Karya, Bali, were most helpful during JFV’s all too brief stay in January 2010 and are

much thanked for lodging, food, materials, transport, and access to their office and herbarium.

Dr. J.C. Regalado Jr. (Hanoi) pointed out Philippine Medinilla species with more-or-less similar

inflorescences. Dr. Irawati (KRB) provided some photographs. The superb drawing was made

by Ms. A. Walsmit Sachs-Jansen, L.

References

Backer, C.A. & Bakhuizen van den Brink f, R.C. (1964) Melastomataceae. Flora of

Java |: 354-374. Groningen: Noordhoff.

Bakhuizen van den Brink Jr., R.C. (1943) A contribution to the knowledge of the

Melastomataceae occurring in the Malay Archipelago, especially in the Netherlands

East Indies. Thesis, Utrecht: 1-391. [For the complicated history of this publication,

_ see Stafleu & Cowan (Taxonomic Literature, ed. 2, 1: 106-107. 1976). This is a

preprint of the next two, which were published using the original type set. |

Bakhuizen van den Brink Jr., R.C. (1946) Rec. Trav. Bot. Néerl. 40: 1-391.

Bakhuizen van den Brink Jr., R.C. (1947) Meded. Bot. Mus. Herb. Rijks Univ. Utrecht

91: 1-391.

Bodegom, S. & Veldkamp, J.F. (2001) A revision of the pseudo-stipular species of

Medinilla Gaud. ex DC. (Melastomataceae - Melastomatoideae - Miconieae).

Blumea 46: 527-567.

Elmer, A.D.E. (1911) New Melastomataceae. Leafl. Philipp. Bot. 4: 1208. Manila:

Elmer.

Gleason, H.A. (1939) The genus Clidemia in Mexico & Central America. Brittonia 3:

97-140.

Hemsley, W.B. (1895) Flora of the Solomon islands. Bull. Misc. Inform. Kew: 132-

Is97@dss):

H6, P.H. (1992) Cay co Viétnam 2, 1: 114, t. 3938. Montréal: Ho.

Merrill, E.D. (1910) New or noteworthy Philippine plants, VII. Philipp. J. Sci., C 5:

DOR

Merrill, E.D. (1913) Studies on Philippine Melastomataceae , I. Philipp. J. Sci., C 8:

235, 247, 248.

Mildbread, J. (1922) Wissenschaftliche Ergebnisse der zweiten Deutschen Zentral-

Afrika-Expedition, 1910-1911, 2: 116. Leipzig: Klinkhardt & Biermann.

Quisumbing, E. & Merrill, E.D. (1928) New Philippine plants. Philipp. J. Sci. 37:

180-181.

Regalado, Jr., J.C. (1990) Revision of Medinilla (Melastomataceae) of Borneo. Blumea

35: 5—70.

Regalado, J.C. (1995) Revision of Philippine Medinilla (Melastomataceae). Blumea

40: 152-153, 181-183, t. 16.

Renner, S.S., Clausing, G., Cellinese, N. & Meyer, K. (2001) Melastomataceae. Flora

of Thailand 7: 412-497. Bangkok: Forest Herbarium.

Robinson, C.B. (1911a) Botanical notes upon the island of Pollilo. Philipp. J. Sci., C

6: 217-218.

Robinson, C.B. (1911b) Alabastra philippinensia, III. Philipp. J. Sci., C 6: 351-352.

- a s

Anew Medinilla from Celebes

Schatz, G.E. & Wendt, T. (2004) A new flagelliferous species of Stenanona

(Annonaceae) from Mexico, with a review of the phenomenon of flagelliflory.

Lundellia 7: 28-38.

Van der Pil, L. (1982) Principles of Dispersal in Higher Plants, ed. 3: 27. Berlin:

Springer-Verlag.

Weberling, F. (Pankhurst, R.J., transl.) (1989) Morphology of Flowers and

Inflorescences. 233-234. Great Britain: Cambridge University Press.

Wu, C.Y. (1979) Medinilla, in: Chen, C. (ed) Flora Yunnanica 2: 133, t. 32, f. 1-5.

Beijing: Science Press.

Gardens’ Bulletin Singapore 62 (2): 223-238. 2011

A natural hybrid between Dendrocalamus pendulus

and Gigantochloa scortechinii

(Poaceae: Bambusoideae: Bambuseae)

in Peninsular Malaysia

W.L. Goh’, S. Chandran', K. Kamiya? and K.M. Wong?

‘Institute of Biological Sciences, Faculty of Science,

University of Malaya, 50603 Kuala Lumpur, Malaysia

weilim_goh@yahoo.com (corresponding author)

* Faculty of Agriculture, Ehime University, 3-5-7 Tarumi Matsuyama Ehime 790-8566, Japan

’ Singapore Botanic Gardens, | Cluny Road, Singapore 259569

wkm2000@gmail.com

ABSTRACT. A natural inter-generic bamboo hybrid between Dendrocalamus pendulus

and Gigantochloa scortechinii is reported for Peninsular Malaysia. The hybrid has some

morphological characteristics of each parent but also shows intermediacy between the

parents. We demonstrate the hybridisation using the partial Granule-Bound Starch Synthase

(GBSS) I gene sequence. The nothogenus * Gigantocalamus K.M.Wong and nothospecies

* Gigantocalamus malpenensis K.M.Wong are proposed for the hybrid. We suggest that

chloroplast introgression could have occurred among the parental taxa or their ancestors and

that the significance of introgressive hybridisation in the complex taxonomic relationships of

woody tropical bamboos in SE Asian Bambuseae has not been fully appreciated.

Keywords. Bambuseae, bamboo, Dendrocalamus, * Gigantocalamus, Gigantochloa,

hybridisation, introgression, Malay Peninsula

Introduction: hybridisation in bamboos

Notions of bamboo hybridisation have not had a very long history. In the Malay

Peninsula and Java, Gigantochloa Kurz shows an interesting diversity that includes

many taxa existing or known only in cultivation, which Holttum (1958) postulated

could have originated from selection of the products of hybrid swarms in ancient times.

The natural distribution of Gigantochloa is believed to be centred in the region from

Myanmar to northern Malay Peninsula, and the historical migration of peoples from

mainland SE Asia southwards could have brought such selected clones to Java and

Peninsular Malaysia. More recently, the observations of Muller (1998) on variation

among the offspring of a cultivated clump of Gigantochloa ridleyi Holttum in his

farm at Mount Mirinjo, Queensland, and originating from Bali, provided very good

corroboration for Holttum’s postulation. At Mount Mirinjo, a single clump of this

species flowered, so the seeds were the products of self-fertilisation. Seed set was very

poor and seedling mortality, including albino forms, was significant. The confounding

diversity in morphology produced among the surviving seedlings and young clumps,

most of which had no close resemblance to the mother clump, was reminiscent of the

multiple assortments that could be produced in the F2 generation from the selfing of a

hybrid (Muller 1998).

224 Gard. Bull. Singapore 62 (2) 2011

Some degree of fertility among different subtropical bamboo taxa has been

demonstrated. Zhang & Chen (1980) have reported a successful artificial hybrid,

‘Cheng Ma Qing No.1’, using the pollen of Dendrocalamus latiflorus Munro and

Bambusa pervariabilis McClure as the seed parent, in experimental crossings. The

hybrid was reported to be fast-growing and having hard culms, highly resistant to cold,

and developing ornamental pale-yellow stripes on the lower internodes, making it a

good cultivar for various uses. It showed intermediacy in morphological characteristics

and chromosome number compared to the parental species (Zhang & Chen 1980,

Zhang 1985). On the basis of chromosome numbers, morphological similarity to an

experimental hybrid B. fextilis McClure = D. Jatiflorus, and a high degree of pollen

sterility, Zhang (1985) also suspected B. stenoaurita (W.T.Lin) T.H.Wen to be a natural

Fl hybrid of the two species. Indeed, B. stenoaurita has proven difficult to classify

and has been transferred from one genus to another without clear resolution (Xia et

al. 2007, Yang et al. 2010). However, no natural bamboo hybrids in tropical Asia have

been clearly demonstrated to date.

Working with temperate bamboos, Muramatsu (1981) suggested that only a weak

crossing barrier exists among woody bamboos, which makes both inter-specific and

inter-generic hybridisation highly possible. He arrived at this idea based on pollination

and germination experiments with Phyllostachys Siebold & Zucc., Pleioblastus Nakai

and Sasa Makino & Shibata. The monotypic Hibanobambusa Maruy. & H.Okamura

was established as a hybrid genus (Maruyama et al. 1979) and proven to have originated

from hybridisation between the distantly related Phyllostachys and Sasa (Takahashi et

al. 1994).

Natural hybridisation among American bamboos was first reported by Clark

et al. (1989). Their study of three natural hybrids, Chusquea subtessellata Hitche.

x C. amistadensis L.G.Clark, Davidse & R.P.Ellis, C. subtessellata x C. vulcanalis

(Soderstr. & S.Calderon) L.G.Clark and C. spencei Ernst < C. tessellata Munro, also

showed intermediate morphological and anatomical characteristics of their respective

parental species. These hybrids displayed normal meiosis stages and high pollen

fertility. Hybridisation was expected to be significant in the evolution of Chusquea

sect. Swallenochloa (McClure) L.G.Clark to which these taxa conform, as some of the

species exhibit a nearly continuous flowering habit, which increases the probability of

hybridisation. Hybridisation was also suggested as a cause of taxonomic difficulties

in the Arundinaria complex (McClure 1973) but this has only been investigated

genetically when Triplett et al. (2010) demonstrated an F1 natural hybrid between A.

gigantea (Walter) Muhl. and A. ftecta (Walter) Muhl. using the Amplified Fragment

Length Polymorphisms (AFLP) technique and cpDNA phylogenetic analysis.

Furthermore, multiple, reciprocal hybridisation and introgression events were

implicated based on the complex mosaic pattern of the genetic composition in the

three hybrid individuals. Their complex origin involves not only 4. gigantea and A.

tecta, but also A. appalachiana Triplett, Weakley & L.G.Clark. It is also noteworthy

that not all of these hybrid individuals showed morphological intermediacy (Triplett

et al. 2010).

The present study provides molecular evidence for an inter-generic bamboo

hybrid that shows a combination of morphological features of its parental species,

Dendrocalamus pendulus Ridl. and Gigantochloa scortechinii Gamble. Considering

that allelic heterozygosity could be a strong indication of an F1 hybrid status, we

sequenced the GBSS/ gene of the putative hybrid individuals and their parental species

Natural hybrid bamboo in Peninsular Malaysia 225

to demonstrate their relationship. This approach is suitable for the current sampling

scale in terms of cost- and time-effectiveness.

Materials and methods

Plant materials

A putative natural hybrid (hereafter referred to as Hybrid Tapah) was collected from

among Dendrocalamus pendulus and Gigantochloa scortechinii clumps along the

Tapah-Cameron Highlands road, Peninsular Malaysia, on 28 November 2001. Material

raised from a rhizome offset was planted in the Bambusetum, Rimba Ilmu Botanical

Garden, University of Malaya. This clone (Fig. 1) flowered in April 2007, i.e., quite

soon after it grew to mature size and then died completely in July 2008. Voucher

material was deposited with the Herbaria of the University of Malaya, Kuala Lumpur

(KLU), Singapore Botanic Gardens (SING) and Iowa State University (ISC), and leaf

material dried in silica gel was obtained for molecular studies.

A population of the same putative hybrid encountered in 2009 along the Old

Gombak Road, Selangor, Peninsular Malaysia, again sympatric with D. pendulus and

G. scortechinii clumps, was also studied (Fig. 2). Voucher material of two individuals

(Hybrid Gombak-1 and Hybrid Gombak-2) was collected and deposited with KLU,

detailed morphological observations were made, and leaf material dried in silica gel

was also obtained.

Besides these three hybrid accessions, three accessions of leaf material of D.

pendulus and five accessions of G. scortechinii were likewise obtained. Identification

followed Wong (1995). Voucher reference numbers and collection localities are shown

in Table 1.

DNA extraction and polymerase chain reaction (PCR)

Total DNA was extracted from silica-dried young leaves using Qiagen DNeasy

Extraction kits following instructions by the manufacturer. Polymerase chain reaction

(PCR) was run using a Perkin Elmer GeneAmp 9600 Thermocycler with the programme

set at 2 min at 95.0°C: 30 cycles of 30 a at 94.0°C, 45 s at annealing temperature, |

min at 72.0°C; 5 min at 72.0°C; hold at 4.0°C. Annealing temperatures were 59.0°C

for primers Gin (forward) and GBSS (reverse) and 55.0 °C for primers for cpDNA,

rps16-trnQ, trnC-rpoB, trnH-psbA and trnD-T (Bamboo Phylogeny Group 2005). The

DNA markers have been useful in resolving the phylogenetic relationships among

some Dendrocalamus and Gigantochioa taxa in the analyses using both GBSS/ and

rps16-trnQ + trnC-rpoB + trnH-psbA + trnD-T (Goh et al. 2010). The PCR reaction

mixture contains 1.5 mM MgCl2, 0.5 uM forward and reverse primers each, 0.2 mM

of dNTPs, 1x PCR buffer and ~10 ng of DNA samples. PCR products were purified

using Promega PCR Clean-up System kits following instructions by the manufacturer.

PCR cloning, haplotype-specific primer design and DNA sequencing

Purified PCR products for the partial GBSS/ gene of the putative hybrid individuals

were ligated into pDrive vectors and transformed into EZ competent cells following

the instructions of the Qiagen PCR Cloning Plus kit. White colonies were picked to

perform colony-PCR using the primers Gin (forward) and GBSS (reverse). Nine to

fifteen clones for each hybrid individual were successfully amplified and sequenced.

Gard. Bull. Singapore 62 (2) 2011

Fig 1. The putative hybrid Dendrocalamus pendulus < Gigantochloa scortechinii. A. Clump

habit. B. Culm shoot. C. Culm internode characteristics. D. Pseudospikelet cluster. Photo

credits: M. Sugumaran.

Natural hybrid bamboo in Peninsular Malaysia 227

Fig 2. Culm shoots of the two parental species and their putative hybrid from clumps within

the hybrid zone along the Gombak road, Selangor, Peninsular Malaysia. A. Gigantochloa

scortechinii. B. Dendrocalamus pendulus. C. Putative hybrid. Photo credits: K.M. Wong.

The sequences of all clones were aligned. Three indel regions and a number of variable

sites were observed in the DNA sequences of the clones. As some of the unique

nucleotide substitutions could be possibly due to PCR or cloning errors, we designed

two sets of internal primers specific for each GBSS/ haplotype in order to obtain

unambiguous DNA sequence for each allele. The location of haplotype-specific primers

are shown in Fig. 3. Internal primers Gin336/1 (forward) and Gin336/2 (forward) were

designed for the indel region, and Gin396/1 (reverse) and Gin396/2 (reverse) were

designed for the region containing three variable sites. Primer sequences are shown

in Table 2. PCR was performed using the following primer-pairs: (1) Gin—Gin396/1,

(11) Gin—Gin396/2, (111) Gin336/1—GBSS, and (iv) Gin336/2—GBSS for each putative

hybrid individual. Direct sequencing of purified PCR products was commercially done

by FirstBase Laboratory Sdn. Bhd. (Malaysia). For each hybrid accession, sequences

generated using the primer-pairs (i) and (111) were merged as a haplotype, and those

generated using the primer-pairs (ii) and (iv) were merged as another haplotype. All

the sequences obtained were deposited in GenBank (Table 1).

DNA sequence analysis

Partial GBSS/ gene sequences of putative hybrid individuals were aligned with those

of the putative parental species using ClustalX (Thompson 1997) and manually

adjusted using Bioedit v7.0.9 (Hall 1999) where necessary. Variable sites and indels

were extracted and tabulated.

Phylogenetic analyses

Sequences representing the putative hybrid, D. pendulus and G. scortechinii were

subjected to maximum parsimony (MP) analysis based on four cpDNA non-coding

regions, rps16-trnQ, trnC-rpoB, trnH-psbA and trnD-T, and the partial nuclear GBSS/

region. Holttumochloa magica (Ridl.) K.M.Wong and Kinabaluchloa nebulosa (Stapf. )

228 Gard. Bull. Singapore 62 (2) 2011

K.M.Wong (GenBank accession numbers given in Table 1) were used as the outgroup

because of their sister relationship to the Bambusa-Dendrocalamus-Gigantochloa

complex (BDG complex; Goh et al. 2010). Dinochloa malayana S.Dransfield

(Genbank accession number given in Table 1), a sister-taxon of the BDG complex-H.

magica-K. nebulosa alliance, was also included for a better polarization between the

ingroup and outgroup. Maximum parsimony analysis was performed using PAUP

4.0 b10 (Swofford 2002). A strict consensus tree was reconstructed using heuristic

Table 1. List of accessions of Dendrocalamus pendulus, Gigantochloa scortechinii, the hybrid,

and outgroup taxa, with collection localities and GenBank accession numbers.

Accession

D. pendulus-1

D. pendulus-2

D. pendulus-3

G. scortechinii-|

G. scortechinii-2

G. scortechinii-3

G. scortechinii-4

G. scortechinii-5

Hybrid Gombak-1

Hybrid Gombak-2

Hybrid Tapah

Dinochloa

malayana (outgroup)

Holttumochloa

magica (Outgroup)

Kinabaluchloa

nebulosa (outgroup)

Collection localities / Literature

reference and voucher numbers

if available

Gombak Road, Selangor, Peninsular

Malaysia

Gombak Road, Selangor, Peninsular

Malaysia

Rimba Ilmu Botanical Garden, Univ.

of Malaya, Peninsular Malaysia /

GWL 6 (KLU)

Hulu Langat, Selangor, Peninsular

Malaysia / GWL 2 (KLU)

Chebar, Kedah, Peninsular Malaysia

Gombak Road, Selangor, Peninsular

Malaysia

Road from Kuala Kubu Baru to

Fraser Hill, Peninsular Malaysia /

GWL 9 (KLU)

Rimba Ilmu Botanical Garden, Univ.

of Malaya, Peninsular Malaysia /

Bambusetum Acc. 52

Gombak Road, Selangor, Peninsular

Malaysia / GWL 13 (KLU)

Gombak Road, Selangor, Peninsular

Malaysia / GWL 14 (KLU)

Tapah-Cameron Highlands road,

Peninsular Malaysia / WKM 2895

(KLU, SING)

Goh et al. (2010)

GenBank accession numbers

rps16-trnQ, trnC-rpoB, trnH-psbA,

trnD-T, GBSSI

HQ697856, HQ697867, HQ697903,

HQ697878, HQ697889

HQ697857, HQ697868, HQ697904,

HQ697879, HQ697888

HQ697855, HQ697866, HQ697902,

HQ697877, HQ697890

HQ697861, HQ697872, HQ697908,

HQ697883, HQ697897

HQ697863, HQ697874, HQ697910,

HQ697885, HQ697898

HQ697864, HQ697875, HQ697911,

HQ697886, HQ697899

HQ697865, HQ697876, HQ697912,

HQ697887, HQ697900

HQ697862, HQ697873, HQ697909,

HQ697884, HQ697901

HQ697860, HQ697871, HQ697907,

HQ697882, (HQ697891,

HQ697894)

HQ697859, HQ697870, HQ697906,

HQ697881, (HQ697892,

HQ697895)

HQ697858, HQ697869, HQ697905,

HQ697880, (HQ697893,

HQ697896)

FJ416343, GU390924, GU390951,

GU390973, GU391005

FJ416348, GU390931, GU391012,

GU390958, GU390980

FJ416356, GU390932, GU391013,

GU390959, GU390981

\Oo

Natural hybrid bamboo in Peninsular Malaysia

search with 10 random sequence additions and TBR branch swapping. ‘MulTrees’

was limited to 10000 trees. Bootstrap analysis was run using 1000 replicates. The

potentially informative indels were scored following the Simple Indel Coding (SIC)

method (Simmons & Ochoterena 2000).

The best model for the Bayesian Inference (BI) analysis was tested using

MrModeltest 2.2 (Nylander 2004). BI analyses were performed in MrBayes 3.1

(Huelsenback & Ronquist 2001), using 2 runs of 4 chains each, and run for 1000000

generations with trees sampled every 100 generations. The first 2500 trees were

discarded as burn-in.

Table 2. Sequences of PCR primers for the partial GBSS/ gene.

Primer Forward/ Sequence (5’ - 3’) Source

Reverse

Gin Forward AAG TTT GAG CGC ATG TTC CAG AGC | Gohetal. (2010)

GBSS Reverse GGC GAG CGG CGC GAT CCC TCG CC Mason-Gamer et

al. (1998)

Gin336/1 Forward CLEMIAN GRC E TECANG CAGE This contribution

Gin336/2 Forward Cie CINGLE iC MG CAG Ere This contribution

Gin396/1 Reverse CAA GAG TAA CGC CAT ATA TG This contribution

Gin396/2 Reverse CAA GAG TAA CAC CAT GTA CG This contribution

Site 1 221 336348 396 609 72)

— —

— > + an

Gin Gin336/1 Gin396/1 GBSS

ae od

Gin336/2 Gin396/2

Fig. 3. Schematic diagram showing the position of haplotype-specific primers (site numbers) and

indel regions (grey bars) in the partial GBSS/ gene. Arrows indicate directions of primers.

Results

Morphological characteristics and reproductive behaviour of the hybrid

A comparison of the morphological characters of the hybrid and its parental species

is shown in Table 3. As has been noted for many hybrids and hybrid derivatives

(Rieseberg 1995), the morphology of the hybrid is a mixture of qualitative characters

that match one or the other parental species, e.g., culm leaf auricles with bristles in

the hybrid and Dendrocalamus pendulus (Fig. 2), and fused staminal filaments in the

hybrid and Gigantochloa scortechinii, or are intermediate between the parents, e.g.,

length of the pseudospikelets. During the entire flowering period of Hybrid Tapah and

up to a month afterwards, no caryopses were found in spite of careful searches.

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Natural hybrid bamboo in Peninsular Malaysia 231

Sequence characteristics of the partial GBSSI gene

The haplotypes of each putative hybrid individual are called Haplotype D and

Haplotype G, respectively. Haplotypes D and G were 705-706 bp in length. Multiple

DNA sequence alignment of GBSS/ haplotypes of the hybrid, D. pendulus and G.

scortechinii revealed that 26 out of 35 variable/ indel sites are indicative of the

parentage of the hybrid (Table 4).

Maximum parsimony (MP) and Bayesian Inference (BI) analyses

The aligned data matrix of the partial GBSS/ gene for the ingroup consists of 707

characters, of which 26 are parsimony-informative. MP analysis resulted in four

equally most parsimonious trees (shown in Fig. 4). Bayesian analysis using Model

K80 generated a similar topology. All five G. scortechinii accessions form a clade with

the G haplotypes of the hybrid accessions, whereas all three D. pendulus accessions

form a clade with the D haplotypes of the hybrid accessions (Fig. 4).

The aligned data matrix of the combined cpDNA, rps/16-trnO + trnC-rpoB +

trnH-psbA + trnD-T dataset for the ingroup consists of 3889 characters, of which 26

are parsimony-informative. MP analysis resulted in four equally most parsimonious

trees (shown in Fig. 5). Bayesian analysis using Model HK Y~+I generated a similar

topology. One of the major clades was formed by all three D. pendulus accessions, all

three hybrid accessions, as well as three G. scortechinii accessions. Hybrid Tapah was

at the basal node of this clade. The remaining two accessions of G. scortechinii form

another clade, sister to the other ingroup cluster.

Table 4. The 28 variable sites and 11 indel sites of the partial GBSS/ gene (722 bp) of the hybrid

and its parental species. Dots indicate identical nucleotides compared to those in the first row.

Dashes indicate the alignment gaps. Twenty six sites characterising the hybrid origin of the

hybrid individuals are highlighted.

i161 7 Bee 38 ee 1h 4A i A eS eee eee sc SSG

1 #86 23 5 8 Bees 6 eee O 1 6 6 2 Pee 466712

Taxon 88 491 2 Bee 5S Bee 6 253648023235 437911

D. penduius-i -BGCTAABPAAG——--—-cCGCGcGCcCAACCEGCETATATTIETGCACE

D. pendulus-2 AB..-.-.T . ae = = Be ae & T . . - @C Cee C Bee

D. penduius—3 AB..--.-R a7 Se T he T.. . BC Bee C Cee

Hybrid Gombak-i (haplotype D) ag..... == es T Sea eet TT. . - @ . Gee Cee

De D) ag..-. -_ =e. Sete T es ee be T-— ..@. Gee Cc See

AB. --T -. ia = = SEO. ee T-—.. 8 - Gees - See

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AG. - . - T Ripe Teme Cll. eee Cc Bree

$= AB. .C . Tf Rie Teme C... 8 . Bee C eee

G. AB.tTt..T eee. T eee es C - 2. eee C Cer

Gc. ABT . . T ieee ST eee Cl. . BB . Be Bee

Cc. AR. . . . T Sees Tee Cl. Tl. Ue. Be C Ree

Discussion

Indels and nucleotide substitutions observed in the partial GBSS/ gene sequences

of Hybrid Gombak-1, Hybrid Gombak-2 and Hybrid Tapah (Table 4) suggest that

haplotype D is derived from Dendrocalamus pendulus and haplotype G is derived

from Gigantochloa scortechinii, as expected. This hypothesis was also supported by

the placement of haplotypes D and G in the GBSS/-based topology, where haplotypes

BY) Gard. Bull. Singapore 62 (2) 2011

D form a single clade with D. pendulus and haplotypes G form a single clade with G.

scortechinii (Fig. 4). From the genotypes of the hybrid and its parental species, the

hybrid is reasonably interpreted as a relatively recent F1 offspring.

Assuming cpDNA is maternally inherited in the bamboos, as reported for many

angiosperm taxa (Corriveau & Coleman 1988), we attempted to deduce the seed

parent of the hybrid from the cpDNA-based topology. However, D. pendulus and G.

scortechinii did not form distinct clades. Rather, one of the two clades consists of

D. pendulus, G. scortechinii and the hybrids, and another clade consists of only G.

scortechinii (Fig. 5). Similar inter- and intra-specific cpDNA variations have been

reported for Quercus L. (Whittemore & Schaal 1991, Petit et al. 1997, Bordac et al.

2000, Petit et al. 2008) and such patterns were attributed to interspecific gene flow

resulting from introgression (Lexer et al. 2006) or shared polymorphism (Muir &

Schl6étterer 2005, 2006). Other studies inferring chloroplast capture based on shared

cpDNA haplotype patterns include those for Saxifragaceae (Soltis et al. 1991,

Okuyama et al. 2005), Pinaceae (Watano et al. 1996, Senjo et al. 1999, Ito et al. 2008),

Phlox L. (Ferguson et al. 2002), Salix L. (Hardig et al. 2000) and Nothofagus Blume

(Acosta & Premoli 2010). Two of these studies demonstrated that this chloroplast

introgression has a strong association with geographic distribution rather than with

taxonomic relationships (Whittermore & Schaal 1991, Acosta & Premoli 2010). It is

Hybrid Gombak-1 (Haplotype G)

0.98 | Hybrid Gombak-2 (Haplotype G)

. scortechinii-3

. scortechinii-4

. scortechinil-2

. scortechinil-1

. scortechinii-5

4.00 Hybrid Tapah (Haplotype G)

D. pendulus-1

D. pendulus-3

1.00 | Hybrid Gombak-1 (Haplotype D)

Hybrid Gombak-2 (Haplotype D)

D. pendulus-2

Hybrid Tapah (Haplotype D)

Dinochloa malayana

Holttumochloa magica

96 Kinabaluchloa nebulosa

—+

Fig. 4. One of the four most parsimonious trees from the maximum parsimony analysis based

on the partial GBSS/ region (Tree length = 67, CI = 0.9254, RI = 0.9655). Posterior probabilities

>().90 are shown above the nodes, bootstrap support values >70% below the nodes. The tree

is drawn to scale, with branch lengths indicating evolutionary distances as number of base

substitutions per site.

Natural hybrid bamboo in Peninsular Malaysia ZI

noteworthy that sharing of chloroplast DNA haplotypes was also observed in two well-

defined North American bamboos, Arundinaria tecta and A. appalachiana (Triplett et

al. 2010). Our cpDNA phylogeny is not feasible for determining the direction of the

cross but implies that ancient chloroplast introgression has been possible between D.

pendulus and G. scortechinii. However, other possible explanations for the cpDNA

phylogenetic tree topology, such as reciprocal crosses followed by introgression,

could not be ruled out. Extensive studies including more accessions of hybrid, more

populations of D. pendulus and G. scortechinii, and perhaps more of their congeners

in Peninsular Malaysia are much needed to address this problem.

Through this study, we confirm that natural inter-generic hybridisation occurs

among Malaysian bamboos and suggest that the role of hybridisation in the evolution

of tropical Asian bamboos could have been underestimated. There is circumstantial

evidence for natural hybridisation in bamboos but well-documented instances are

scarce. Some taxa appear to have a high degree of morphological variation resulting in

poorer distinction among species, e.g., the Malayan-Javan Gigantochloa taxa (Holttum

1958). A number of taxa are cultivated for their usefulness but have never been found

in the wild, e.g., the Malaysian village bamboos Bambusa heterostachya (Munro)

D. pendulus-1

D. pendulus-2

Hybrid Gombak-1

G. scortechinii-2

D. pendulus-3

Hybrid Gombak-2

G. scortechinii-3

G. scortechinii-4

Hybrid Tapah

G. scortechinii-1

G. scortechinii-5

D. malayana

H. magica

100 K. nebulosa

Fig. 5. One of the four most parsimonious trees from the analysis using Bayesian inference

based on 4 cpDNA intergenic spacers, rps16-trnQ, trnC-rpoB, trnH-psbA, and trnD-T (Tree

length = 78, CI = 0.9231, RI = 0.8868). Posterior probabilities >0.90 are shown above the

nodes, bootstrap support values >70% below the nodes. The tree is drawn to scale, with branch

lengths indicating evolutionary distances as number of base substitutions per site.

234 Gard. Bull. Singapore 62 (2) 2011

Holttum, B. Jaxa K.M.Wong, D. asper Backer and G. thoii K.M.Wong (Wong 1995,

2004), and many such taxa are infertile or have low fertility (Wong 1995, Ramanayake

1998, Muller 1998, Koshy & Jee 2001), as is the case with our Hybrid Tapah here. In

B. vulgaris Schrad. (Koshy & Jee 2001), seed set failure is due to poor stigma exposure

and pollen sterility resulting from meiotic irregularities; in addition, chromosomal

inconstancy (mosaicism) is also found among root-tip (1.e., somatic) cells. Selfing in a

reproductively isolated G. rid/evi clump had overwhelmingly high seed failure and the

offspring that did survive were morphologically inconsistent with one another and the

parent clump (Muller 1998). We suggest that introgressive hybridisation 1s an important

source of variability and, in view of the demonstrated possibilities for hybridisation

even among genera (Zhang & Chen 1980, Takahashi et al. 1994, and this study), has

contributed to taxonomic complexity in the Bambusinae, particularly the closely related

Bambusa-Dendrocalamus-Gigantochloa complex. A recent phylogenetic study of SE

Asian bamboos (Goh et al. 2010) recovered incongruent elements between nuclear and

chloroplast sequence-based topologies which implied that the possibility of reticulate

evolution in the Bambusinae should be further investigated.

Naming and description of the new hybrid

The detected hybrid has not been previously described or named. It has been produced

by hybridisation between Gigantochloa scortechinii and Dendrocalamus pendulus,

both common bamboos in the foothills of the Main Range in Peninsular Malaysia, in

at least two localities (Tapah and Hulu Gombak) and seems likely to occur in more

localities. Although there has been a suggestion that Dendrocalamus and Gigantochloa

may not be generically distinct (Soderstrom & Ellis 1987), yet currently available

analyses using larger data sets (Yang et al. 2008;, Yang et al. 2010, Goh et al. 2010) have

not clearly resolved this matter. On the other hand, the type species or type alliances

of these genera have been recovered as distinct clades in some analyses (Goh et al.

2010), and from a morphological standpoint, these genera do at least seem to form

very different extremes (as represented by their type species and other closely related

species) (Holttum 1958, Wong 1995). Wong (1995, 2004) notes that there are species

currently placed in Dendrocalamus that vary significantly from the type alliance that

includes D. pendulus.

Here we formally name the nothogenus and nothospecies, according to

the International Code of Botanical Nomenclature (McNeill et al. 2006). As the

nothogeneric name should be formed from combining elements of the names of both

parent genera, and Dendrochloa has been pre-empted by Dendrochloa C.E.Parkinson,

we have opted for the obvious choice, Gigantocalamus. The species epithet is an

abbreviation of “Malay Peninsula”, where this hybrid was first noted.

< Gigantocalamus K.M.Wong, nothogenus nova

[= Dendrocalamus Nees * Gigantochloa Kurz]

Bambusa erecta caespitosa, culmi foliorum vagina abaxialiter parum ad moderate

pallide cerea, in quoque nodo ramus primarius dominans rigidus adscendens distaliter

cernuus, longitudine pseudospicularum inter parentes intermedia, ut videtur seminum

absentia sterilis.

Natural hybrid bamboo in Peninsular Malaysia 235

<x Gigantocalamus malpenensis K.M.Wong, nothospecies nova

[= Dendrocalamus pendulus Ridl. x Gigantochloa scortechinii Gamble]

Hybrida_ naturalis Gigantochloae scortechinii similis, culmi foliorum vaginae

marginibus papyraceis, auriculis lobiformibus rotundatis marginibus setosis,

lemmatibus glabris differt; Dendrocalami penduli similis, culmi foliorum vagina atro-

pilosa, filamentis stamineis tubo connatis differt; in characteribus ceteris inter ambo

species intermedia.

TYPE: Peninsular Malaysia, Kuala Lumpur, University of Malaya, Rimba Ilmu

Botanic Garden, Bambusetum accession no. 48, 13 Apr 2007 (originally collected

from Perak, Tapah-Cameron Highlands road, 28 Nov 2001), Wong et al. WKM 2895

(holo KLU; iso ISC, SING).

Clumped bamboo; culms to c. 10 m tall, at first erect, then arching outwards, diameter

c. 4 cm, internodes 20-25 cm at midculm, green except for pale yellow-green stripes

at the culm base, with a ring of pale matted hairs above and below each node, dark

brown hairs scattered all over, not to only slightly white-waxy. Culm leaf sheath pale

yellow-orange with a slight tint of pink or dark purple-brown, dark brown hairy and

slightly to moderately white-waxy on the back; edge of the sheath very thin, drying as

a thin papery pale-brown marginal zone; auricles rounded lobes to c. 5 mm high, dark

purplish black, with pale stiff wavy bristles to over 10 mm long; blade medium green

and leaflike with a pink flush, spreading to reflexed. Branch complement developing

from a single bud, with one dominant primary central axis and a few smaller higher-

order branches from its base, all branches ascending in habit, the dominant primary

axis tending to droop at its tip. Branch leaf blades pale short-hairy on the lower side.

Pseudospikelets 7—1\1 mm long, developing into clusters of few to many; with (from

the base) 2-4 bracts subtending prophyllated buds, 2—3 empty glumes, 2 (exceptionally

3) florets and a terminal empty lemma (this terminal empty lemma sometimes replaced

by a third floret). Lemmas 6—9 mm long, with a short terminal cusp, stiff and scale-

like, glabrous, green to pink-flushed; paleas 5.5—8 mm long, 2-keeled, membraneous,

glabrous except minutely hairy keels, 5-veined on the back, 1-veined between keel and

margin; stamens 6, staminal filaments fused into a tube and extruded from the lemma

when mature; anthers 3—S mm long with an apical cusp 0.3—0.5 mm long, flushed

pink to pale lilac when fresh, pale when dry; empty; ovary rounded, c. 0.5 mm long,

long-hairy at the summit; style short-hairy throughout, terminating in a single plumose

stigma. Caryopsis unknown, apparently not or rarely formed. (Fig. 1 A—D, 2C)

Distribution: Peninsular Malaysia, along the Tapah-Cameron Highlands road in Perak

(Wong et al. WKM 2895) and along the old Gombak road in Selangor (Goh et al.

GWL13, 14).

ACKNOWLEDGEMENTS. WLG received University of Malaya grants FS275/2008C and

PS151/2008B, and a University of Malaya postgraduate fellowship that enabled her work.

KMW acknowledges the support of IPGRI (now Bioversity International), L.T. Hong and the

late Baya Busu during a field excursion when the hybrid was first discovered along the Tapah-

Cameron Highlands road in Peninsular Malaysia. We thank Dr. J.P. Veldkamp (Nationaal

Herbarium Nederland, Leiden University) for help with the Latin used, and Prof. Xia

236 Gard. Bull. Singapore 62 (2) 2011

Nianhe (South China Institute of Botany) for providing literature on the Guangzhou bamboo

hybridisation experiments and arranging for KMW to view the living hybrid clumps at the

Guangdong Forest Research Institute. Prof. Lynn Clark (lowa State University) and Prof. Xia

have given useful comments on this work.

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Gardens ’ Bulletin Singapore 62 (2): 239-251. 2011 239

A revision of Begonia sect. Sphenanthera (Hassk.) Warb.

(Begoniaceae) from Sumatra

Mark Hughes’ and Deden Girmansyah-

‘Royal Botanic Garden Edinburgh, 20a Inverleith Row, Edinburgh, EH3 SLR, U.K.

m.hughes@rbge.ac.uk (corresponding author)

* Herbarium Bogoriense, Cibinong Science Center, Java, Indonesia

ABSTRACT. Begonia sect. Sphenanthera is characterised by robust herbs with fleshy fruits.

Three species are recognised from Sumatra (Begonia longifolia Blume, B. multangula Blume

and B. scottii Tebbitt) and three names have been reduced to synonyms (B. sarcocarpa Ridl.

and B. turbinata Ridl.= B. longifolia Blume and B. trigonocarpa Ridl. = B. multangula Blume).

All species are considered to belong to the IUCN category Least Concern as they have wide

distributions within Sumatra and are known from several localities.

Keywords. Begonia sect. Sphenanthera, conservation status, Sumatra, taxonomy

Introduction

With a rare exception (Kiew 2005) modern monographs and revisions of Southeast

Asian Begonia are notable for their absence. Reasons for this include the large number

of species, a high proportion of undescribed taxa, uncertainty regarding sectional

classification and the fact that Begonia tend to make poor herbarium specimens.

Recently, baseline information regarding sectional placement (Doorenbos et al.

1998) and regional species lists (Hughes, 2008) has become available for facilitating

revisionary work. In order to make the task more manageable still, it has been decided

to tackle the considerable Begonia diversity of Southeast Asia through a series of

regional sectional revisions, beginning with Sumatra and Begonia sect. Sphenanthera.

Begonia sect. Sphenanthera currently contains 17 accepted names from Asia.

Most of the species diversity is on the continent, tapering off towards the east of the

Sunda Shelf and only two species occur east of Wallace’s line. The section contains

quite robust species, which are most obviously characterised by bearing fleshy fruit,

and also flowers with often quite fleshy tepals and male flowers with large androecia

consisting of stamens with extended connectives. It bears a strong resemblance to

Begonia sect. Platycentrum (which differs in having 2-locular dehiscent fruits), and

indeed molecular evidence shows Begonia sect. Sphenanthera to be polyphyletic

and nested within it (Tebbitt et al. 2006). Nevertheless, Begonia sect. Sphenanthera

continues to be a useful taxonomic unit in partitioning the otherwise unwieldy diversity

of the genus, and its status is upheld here.

The revision is based on consulting specimens of Begonia sect. Sphenanthera

from the herbaria A, ANDA, BM, BO, E, FI, K, L, SING and field work in Aceh, North

Sumatra and West Sumatra during 2007—2009. Species classified within Begonia sect.

Sphenanthera tend to be amongst the most widespread in the genus, presumably

because of the fleshy berry-like fruit syndrome, an adaptation to animal dispersal of

their seeds. The combination of being widespread and having a reasonable amount of

240 Gard. Bull. Singapore 62 (2) 2011

population variability, plus the tendency to form hybrids, means the level of synonymy

and taxonomic complexity in species from Begonia sect. Sphenanthera is higher than

in other Asian sections of the genus. It also means a wider species concept needs to be

applied, particularly as the propensity for forming narrow endemics is not as high as

for other Asian sections. Three species are recognised from Sumatra in this revision,

and three species are reduced to synonyms. Only one of the species, Begonia scottii,

is endemic to Sumatra.

Currently, 52 Begonia species are recognized from Sumatra (Hughes 2008,

Hughes et al. 2009), with most occurring along the Barisan range, which forms the

spine of much of the island and in the mountains of North Sumatra and Aceh. Although

there may be some further synonymy to be uncovered, the total number of species, and

of endemic species, is likely to rise significantly.

Taxonomic part

Begonia sect. Sphenanthera (Hassk.) Warb., Nat. Pflanzenfam. 3(6A): 141 (1894). —

Sphenanthera Hassk., Versl. Akad. Wetensch. Amsterdam, iv: 139 (1855) — Casparya

sect. Sphenanthera (Hassk.) A.DC., Ann. Sci. Nat. Bot., [IV 11: 118 (1859). TYPE:

Begonia robusta Blume.

Casparya sect. Holoclinium A.DC., Ann. Sci. Nat. Bot. 4(11): 118 (1859). — Begonia

sect. Holoclinium (A.DC.) Warb., Nat. Pflanzenfam. 3(6A): 142 (1894). TYPE:

Casparya trisulcata A.DC. (= Begonia longifolia Blume).

Casparya sect. Polyschisma A.DC., Ann. Sci. Nat. Bot. 4(11): 119 (1859). — Begonia

sect. Polyschisma (A.DC.) Warb., Nat. Pflanzenfam. 3(6A): 149 (1894). TYPE:

Casparya crassicaulis A.DC. (= Begonia multangula Blume)

Robust erect or sprawling fleshy herbs. Stems not woody, stipules large and keeled.

Leaves asymmetric. Inflorescences bisexual; male flowers with a large androecium

giving rise to buds with a distinct bulge, anthers subequal, dehiscing laterally along

the length of the anther, connectives projecting; female flowers with 3 greenish-yellow

large stigmas, spirally or convolutely twisted. Fruit fleshy, indehiscent, three locular,

each locule with two placentae.

Key to species of Begonia sect. Sphenanthera from Sumatra

1. Leaves sub-orbicular to broadly ovate, with S—7 broad pointed lobes ............:ceee

vsouniva ne ddh dence eu horas ant ta oNeGaH oat sec eangeE REN See cena ta eee eee 2. Begonia multangula

2... Leaves ovate truits Dorie in parson TOUS pe. eee eee 3. Begonia scottii

Leaves lanceolate, fruits borne in clusters of 2—15 ............. 1. Begonia longifolia

Begonia sect. Sphenanthera from Sumatra 241

1. Begonia longifolia Blume, Catalogus 102 (1823); Blume, Enum. Pl. Javae 1: 97

(1827); Candolle, Prodr. 15(1): 398 (1864); Koorders, Exkurs.—Fl. Java 650 (1912);

Tebbitt, Brittonia 55: 25 (2003); Tebbitt, Begonias 168 (2005); Kiew, Begonias

Penins. Malaysia 107 (2005); Gu, Peng & Turland, Fl. China 13: 184 (2007); Hughes,

An Annotated Checklist of Southeast Asian Begonia 72 (2008). — Diploclinium

longifolium (Blume) Miq., Fl. Ned. Ind. 1(1): 687 (1856). TYPE: Java, Salak, C.L.v.

Blume 740 (holo B).

Casparya trisulcata A.DC., Ann. Sci. Nat. Bot., [V 11: 119 (1859); Candolle, Prodr.

15(1): 277 (1864). — Begonia trisulcata (A.DC.) Warb., Nat. Pflanzenfam. 3(6A): 142

(1894). TYPE: Java, Mt. Jojing, 1 May 1845, H. Zollinger 2850 (holo G-DC; iso B,

BM, P [2 sheets]).

Begonia inflata C.B.Clarke in Hook.f., Fl. Brit. Ind. 2: 636 (1879); Clarke, J. Linn.

Soc., Bot. 18: 115 (1881); Burkill, Rec. Bot. Surv. India 10: 412 (1924); Craib, FI.

Siam 1: 774 (1931); Grierson, Fl. Bhutan 2: 242 (1991); Tebbitt, Brittonia 55(1): 25

(2003); Uddin, J. Econ. Tax. Bot. 31(3): 594 (2007). TYPE: W. Griffith 2587 (lecto K:

isolecto B, GH n.v., K, P here designated).

Begonia sarcocarpa Ridl., J. Fed. Malay States Mus. 8(4): 38 (1917) syn. nov.; Tebbitt,

Brittonia 55: 27 (2003). TYPE: Sumatra, West Sumatra, Korinchi, Barong Baru, 5 Jun

1914, H.C. Robinson & C.B. Kloss 61 (holo BM).

Begonia tricornis Ridl., J. Roy. Asiat. Soc., Straits Br. 75: 35 (1917); Tebbitt, Brittonia

55(1): 25 (2003). TYPE: Peninsular Malaysia, Pahang, Telom, Nov 1900, H.N. Ridley

14123 (holo SING; iso K).

Begonia turbinata Ridl., J. Fed. Malay States Mus. 8(4): 37 (1917) syn. nov.; Tebbitt,

Brittonia 55: 28 (2003). TYPE: Sumatra, West Sumatra, Korinchi, Siolak Dras, 15

Mar 1914, H.C. Robinson & C.B. Kloss s.n. (lecto BM; isolecto BM, K designated by

Tebbitt 2003 Joc. cit.).

Begonia crassirostris Irmsch., Mitt. Inst. Allg. Bot. Hamburg 10: 513 (1939); Tebbitt,

Brittonia 55(1): 25 (2003). TYPE: Hainan, Lam Ko District, Lin Fa Shan, 2 Aug 1927,

Tsang Wai Tak 278 (lecto E; isolecto Gn.v., Kn.v., MO n.v., UC n.v. here designated).

Begonia roxburghii auct. non (Mig.) A.DC. sensu Ridley J. Fed. Malay States Mus. 4:

20 (1909), Fl. Malay Penins. 1: 854 (1922).

Plant erect, 40—100(—200) cm high. Stem up to 2 cm thick at the base, c. 5 mm thick

towards the apex, not woody, green to red, internodes around 8—12 cm apart, glabrous

or with minute glandular hairs. Stipules deciduous, lanceolate, keeled, glabrous, c. 25

x 7mm, tip extended. Leaves: petiole 2—7 cm long; lamina lanceolate, base shallowly

cordate, strongly asymmetric, upper surface dark green, (6—)10—18 = 2.5—6(—10)

cm, midrib (5—)8.5—14 cm, underside pale green, glabrous or with scattered short

hairs on the veins, upper surface darker green, usually glabrous or sometimes with

scattered short bristly hairs, venation pinnate-palmate; margin dentate-denticulate;

apex acute. Inflorescences axillary, cymose, branching |—3 times, bisexual, male and

242 Gard. Bull. Singapore 62 (2) 2011

female flowers open at the same time; bracts caducous, lanceolate, 6—12 x 2-4 mm;

primary and secondary peduncles stout, distinctly swollen near the nodes, 2 mm thick,

primary c. | cm long, secondary shorter, each unit consisting of | male and 2 female

flowers, (3—)7—15 flowers in total, bracteoles often present on terminal female flowers,

bracteoles 5 x 1-2 mm. Male flowers: pedicels 25-30 mm; tepals 4, white; outer 2

elliptic-orbicular, slightly cup-shaped, fleshy, with a thinner rim, 10-12 « 8-9 mm;

inner 2 spathulate-elliptic, 9-10 x 6-7 mm; androecium symmetric, a loose globose

cluster; stamens bright yellow, 35—90, subequal; filaments slightly shorter than the

anthers, more so for the outer stamens; anthers c. 2 mm long, linear with a rounded tip,

dehiscing through lateral slits running almost the entire length of the anther. Female

flowers: pedicels 7-12 mm; ovary pale green, fleshy, lobed-triangular in cross-section,

c. 7-9 x 10 mm, with three ridges or small fleshy wings on the lobes, 3-locular, placentae

bifid; tepals (rarely 4—)6, white, elliptic-spathulate, subequal, 8-14 x 5—7 mm; styles

3, greenish yellow, deciduous, bifid, c. 4 mm long, papillose surface spirally twisted.

Fruit borne in clusters of (1—)2—15, green, fleshy, globose-triangular, pendulous on a

stiff pedicel, 10-14 mm diameter, apex sometimes slightly extended. (Fig. 1)

Fig. 1. Begonia longifolia habit (main photo), female and male flowers (inset top left) and

ripening fruit (inset bottom right). All from one plant, Gunung Sorik Merapi, North Sumatra.

Photo credits: Mark Hughes.

Material seen: SUMATRA. Aceh: Gunung Leuser Nature Reserve, Air Panas, 19 Mar

2008, P. Wilkie, et al. PW784 (BO, E). North Sumatra: Asahan, Aek Si Tamboerak,

28 Oct 1936, Rhamat si Boeea 10653 (A); Asahan, Dolok Si Manoek-manoek, 5 Oct

1936 - 20 Nov 1936, Rhamat si Boeea 10246 (A, K, L); Asahan, Hoeta Bagasan, 7

Sep 1934 - 4 Feb 1935, Rahmat si Boeea 1082 (A); Baboeli - Paekas, 9 Jan 1932,

Bangham 776 (A); Berastagi Woods, 8 Feb 1921, H.N. Ridley s.n. (BM, K); ibid,

10 Jun 1928, C. Hanel & Rahmat si Boeea 580 (A); Berastagi Woods, West Hill, 14

Feb 1921, H.N. Ridley s.n. (K); Dolok Sibual Buali, 15 Jan 2000, S.J. Davies & S.K.

Begonia sect. Sphenanthera from Sumatra 243

Rambe 2000-44 (A); Gunung Sibayak, 15 Feb 1932, Bangham 1018 (A. K): ibid, 12

May 2007, M. Hughes & D. Girmansyah MH1387 (E [2 sheets]); Gunung Sinabung,

14 May 2007, M. Hughes & D. Girmansyah MH1395 (E); Gunung Sorik Merapi, 17

May 2007, M. Hughes & D. Girmansyah MH1399 (E):; Karoland, 4 Jun 1918, H.H.

Bartlett & C.D. La Rue 100 (L). West Sumatra: Andalas Baruh Bukit, 19 Mar 1986.

Anda collectors 2366 (ANDA); ibid, 19 Mar 1986, Anda collectors 2367 (ANDA);

Barisan Range, Air Sirah, 4 Feb 1981, Anda collectors 208 (ANDA); Bukit Batabuah,

29 Nov 1997, Anda collectors 32 (ANDA): Bukit Kayo Aro, 18 Mar 1999, Anda

collectors 993258 (ANDA); Bukit Tamasoe, Gunung Talang, 28 Jul 1984, M. Hotta

& et al. 197 (ANDA); Desa Ulu Tambulun, 7 Mar 1999, Anda collectors 993228

(ANDA); Gunung Merapi, 29 Nov 1989, Anda collectors 04 (ANDA); ibid, 29 Nov

1989, Anda collectors 07 (ANDA); ibid, 26 Apr 1991, Anda collectors 13 (ANDA);:

ibid, 26 Jul 2009, M. Hughes & A. Taufig MH1572 (ANDA, BO, E): ibid, 19 Jul 2006,

D. Girmansyah, A. Poulsen, I. Hatta & R. Nelvita 759 (E); Gunung Talang. Kayujao,

3 Feb 1989, H. Nagamasu 3546 (ANDA, BO, L): Kerinci-Seblat National Park, Bukit

Sako, 3 May 1996, Anda collectors 7750 (ANDA): Korinchi, Sungei Karing, 2 Mar

1954, 4.H.G. Alston 14040 (BM, L); Korinchi, Sungei Penoh - Indrapura, 8 Mar 1954,

A.H.G. Alston 14312 (A, BM, L); Padang, Limau Manis, 7 Mar 1954, 4.H.G. Alston

14310 (BM); Padang, Lubuk Sulasi, 30 Jun 1953, Jv. Borssum Waalkes 2765 (BO.

L); Pajakumbuh, Mt. Sago, 11 Mar 1989, Anda collectors 10 (ANDA); ibid, 11 Mar

1989, Anda collectors 124 (ANDA); (ANDA); ibid, 5 May 1957, W. Meijer 5752 (L);

ibid, 30 Dec 1907, E. Meijer Drees 7446 (L); Panti Cermin Nature Reserve, | Jun

2007, M. Hughes & D. Girmansyah MH1431 (BO, E); ibid, 2 Jun 2007, M. Hughes

& D. Girmansyah MH1438 (BO, E); ibid, 2 Jun 2007, M@. Hughes & D. Girmansyah

MH1439 (BO, E): Puncak Pato, 12 Mar 1989, Anda collectors 17 (ANDA); Rintis,

11 Sep 1941, A. Surbeck 536 (L); Taman Hutan Raya, Ladang Padi, 27 Jul 2009, M.

Hughes & Nurainas MH1580 (ANDA, BO, E). Bengkulu: 14 Jan 1931, C_N.A. de

Voogd 572 (BO, L).

Notes. Although Ridley transformed our understanding of the Malesian flora, it is

tempting at times to accuse him of being slightly profligate with plant names. Begonia

turbinata Ridl. was upheld by Tebbitt (2003) due to its turbinate fruit and slender,

red-tinged stems. However this species is impossible to separate satisfactorily in the

herbarium, and duplicates were seen carrying conflicting determinations. Also, red-

tinged stems have also been noted in plants with robust stems, and the fruit shape on

the type and isotypes of B. turbinata fit within the range observed in B. longifolia.

Begonia sarcocarpa Ridl. also has the same fruit type as B. /ongifolia, but differs

in its smaller size and having 5 (not 6) tepals in the female flower. However, the

large number of specimens housed at ANDA from West Sumatra show a range of

intermediate sizes linking B. Jongifolia and B. sarcocarpa, and observations in the

field and from cultivated material show that tepal number is not a stable character and

can vary within individuals. Also, B. sarcocarpa is known only from the type, which

has only one or two female flowers. It seems appropriate to consider these two species

as representing population variants of Begonia longifolia.

Ecology and distribution. Bhutan, northeast India, southern China, Burma, Taiwan,

Thailand, Vietnam, Peninsular Malaysia, Sumatra, Java, Lesser Sunda Islands. Within

Sumatra, Begonia longifolia occurs along the whole of the Barisan range at altitudes of

244 Gard. Bull. Singapore 62 (2) 2011

500-1500 m, though most collections are from 900—1200m (Fig. 2). On steep banks,

often near streams, in primary or secondary forest. IUCN category Least Concern.

*s Ye

Sumatera Utara « | Sumatera Utara « Sumbaigin User

E o | oS 3 o

ay | oa 4 Xe,

Ria | Riau % Riau 3

1 r ‘

Sumatera Barat | Sumatera Barat Sumatera Barat

Jambi | 7. Jambi PA

z | : .

Begonia longil a | Begonia muttanguia ¥ ‘ Begon MY

Sumatera Selatan | A Summaiars Scistan ee

Bengkulu ju Bengkulu

| x . oe

4s “sf . |

Lampung | Lampung faced

i | a

re mai = 1 4

Fig. 2. Distribution of Begonia longifolia (left), B. multangula (middle) and B. scottii (right)

on Sumatra.

2. Begonia multangula Blume, Enum. PI. Javae 1: 96 (1827); Candolle, Prodr. 15(1):

275 (1864); Koorders, Exkurs.—Fl. Java 646 (1912); Doorenbos, Begonian 47: 213

(1980); Tebbitt, Begonias 205 (2005); Hughes, An Annotated Checklist of Southeast

Asian Begonia 86 (2008). — Platyvcentrum multangulum (Blume) Migq., Fl. Ned. Ind.

1(1): 695 (1856). — Sphenanthera multangula (Blume) Klotzsch, Bot. Zeitung 15:

181 (1857). — Casparya multangula (Blume) A.DC., Prodr. 15(1): 275 (1864). TYPE:

Java, Blume s.n. (lecto BO, sheet no BO:1818963, here designated).

Platycentrum multangulum var. glabrata (Mig.) Miq., Fl. Ned. Ind. 1(1): 695 (1856);

Klotzsch, Bot. Zeitung 15: 182 (1857). — Begonia multangula var. glabrata Miq.,

Pl. Jungh. 4: 418 (1857). — Casparya multangula var. glabrata (Migq.) A.DC., Prodr.

15(1): 276 (1864). TYPE: Java, Gunung Merapi, FW. Junghuhn s.n. (not located).

Casparya crassicaulis A.DC., Ann. Sci. Nat. Bot., [V 11: 119 (1859); Candolle, Prodr.

15(1): 278 (1864). — Begonia crassicaulis (A.DC.) Warb., Nat. Pflanzenfam. 3(6A):

149 (1894); Smith & Wasshausen, Phytologia 52: 442 (1983). — Begonia pachyrhachis

L.B.Sm. & Wassh., Phytologia 52: 442 (1983). TYPE: WH. de Vriese (holo K n.¥.).

Casparya robusta var. glabriuscula A.DC., Prodr. 15(1): 275 (1864). — TYPE: Java, 1

May 1845, H. Zollinger 2844 (lecto B; isolecto BM, P designated by Hughes (2008)

loc. Cit.).

Begonia trigonocarpa Ridl., J. Fed. Malay States Mus. 8(4): 38 (1917) syn. nov. TYPE:

Sumatra, West Sumatra, Korinchi, Sungei Kumbang, | Apr 1914, H.C. Robinson &

C.B. Kloss s.n. (lecto BM [barcode 00001733]; isolecto BM, K here designated).

Begonia discolor auct. non R.Br. sensu Blume Enum. PI. Javae 1: 96 (1827).

Plant repent to erect, 50-150 cm tall. Stem rhizomatous at the very base, thick, fleshy,

erect portion with internodes 5—15 cm apart, glabrous or with scattered hairs to densely

hairy with long usually white hairs, rarely reddish. Stipules large, semi persistent,

Begonia sect. Sphenanthera from Sumatra 245

broadly lanceolate, keeled, glabrous or hairy along the keel, 40 x 15 mm. Leaves:

petiole 17—25 cm long, glabrous or with short or long hairs, hairs usually white, rarely

reddish; lamina broadly ovate to suborbicular with 5—7 broadly triangular lobes, base

cordate with lobes overlapping, asymmetric. (15—)24-35 = (15—)20-30 cm, midrib

(10—)16—22 cm, venation palmate, margin dentate-denticulate: upper surface glabrous

or with scattered short hairs, lower surface glabrous or with scattered short hairs.

denser on the veins. Inflorescences a cluster of flowers on a long peduncle, axillary,

sometimes appearing pseudo-terminal between a pair of leaves. cymose, branching

twice, male and female flowers open at the same time: bracts caducous, elliptic, c.

1.5 cm long, glabrous or slightly fimbriate; primary peduncle 2.5—-12 cm <x 4-5 mm,

glabrous or with long hairs, secondary peduncles 1—1.5(—3.5) cm, tertiary 34(—10)

mm, remainder of the peduncles very compressed and appearing sub-umbellate.

Male flowers: pedicels c. 15—25 mm long, glabrous or with scattered hairs; tepals

4, white or pale pink; outer tepals orbicular-ovate 20-25 = c. 20 mm, with hairs on

the reverse; inner tepals, orbicular-elliptic, glabrous, c. 15-20 x 15 mm: androecium

symmetric, a loose globose cluster: stamens bright yellow, c. 80 in number: filaments

unequal, 1.5—3 mm long, shorter on the basal stamens: anthers elliptic, c. 1.5 mm long.

connective slightly extended, dehiscing through slits running nearly the entire length

of the anther. Female flowers: pedicels c. 10 mm long: ovary fleshy, lobed-triangular

in cross-section, c. 8-12 mm diameter, with three ridges or small warty wings on the

lobes, 1 wing sometimes larger, 3-locular, placentae bifid: tepals S—6, white, elliptic-

spathulate, subequal, c. 18—20 x 11-14 mm: styles 3, greenish yellow, semi-persistent,

bifid, c. 4 mm long, papillose surface spirally and convolutely twisted. Fruit green

or red, fleshy, globose-triangular, c. 15—18 mm diameter, borne in bunches of around

10—15, with three fleshy ridges, one sometimes more prominent and often slightly

warty. (Fig. 3)

Material seen. SUMATRA. North Sumatra: Gunong Batu Lopang, de Wilde & de

Wilde-Duyfies 1353] (L). West Sumatra: Gunung Merapi, 19 Jul 2006, D. Girmansyah,

et al. 760 (E); Kerinci-Seblat National Park, Bukit Sako, 25 Jan 1995, Anda collectors

2679 (ANDA): ibid, 25 Jan 1985, D. Arbain 4279 (ANDA); Korinchi, Barong Baru, 8

Jun 1914, H.C. Robinson & C.B. Kloss s.n. (BM): Mt. Singalan, O. Beccari HB4513

(FI [2 sheets]); ibid, O. Beccari HB4513A (FI [2 sheets]); Palupuh, 17 Mar 1988,

Anda collectors 166 (ANDA [2 sheets]); Panti Cermin Nature Reserve, 1 Jun 2007,

M. Hughes & D. Girmansyah MH1428B (BO, E); ibid, 1 Jun 2007, M. Hughes &

D. Girmansyah MH1430B (BO, E): ibid, 2 Jun 2007, M@. Hughes & D. Girmansyah

MH1436 (BO, E). Jambi: Korinchi, 21 Jan 2000, S.J. Davies & S.K. Rambe 2000-128

(A): Korinchi, Sungei Penoh, 3 Jun 2004, D. Girmansyah 384 (BO [3 sheets]).

Notes. Begonia multangula is a new record for Sumatra, and this extension of its range

westwards means that it is considerably more variable than previously circumscribed.

Begonia trigonocarpa is reduced here as it fits within the variation of B. multangula

comfortably. The small fruit size as noted by Ridley on the type of B. trigonocarpa

is due to immaturity - the plant is still in flower, and similar size fruits are found on

specimens of B. multangula at the same stage of development. Many of the leaves

on parts of the type specimen are more elongate than is typical for B. multangula on

Sumatra, and in combination with the plant being glabrous this hints at the possibility

of introgression with B. /ongifolia. A small number of individuals on Sumatra have

246 Gard. Bull. Singapore 62 (2) 2011

reddish hairs, and these are probably hybrids with B. scoftii (see notes under that

species) rather than records of the allied B. robusta, which is endemic to western Java.

Begonia multangula is also allied to B. chlorocarpa from Borneo, from which it differs

most noticeably in not having spurred fruit.

Ecology and distribution. Sumatra, Java, Lesser Sunda Islands. Within Sumatra,

Begonia multangula is found in the mountains of the West and North at altitudes

of 700-2500 m, with most collections being from around 1200-1600 m (Fig. 2).

Sprawling on stream banks and slopes in rain forest. IUCN category Least Concern.

Fig. 3. Begonia multangula habit (main photo), inflorescence (inset top left; Gunung Merapi,

West Sumatra) and fruits (insets, bottom). Habit and fruit photographs from Pantai Cermin,

West Sumatra. Photo credits: Mark Hughes and Deden Girmansyah.

3. Begonia scottii Tebbitt, Blumea 50(1): 154 (2005); Hughes, An Annotated Checklist

of Southeast Asian Begonia 72 (2008). TYPE: Sumatra, Aceh, Gunung Leuser Nature

Reserve, Gunung Ketambe, de Wilde & de Wilde-Duyfies 14309 (holo L n.v.; iso BO,

1b)

Plant repent to erect, erect portion 25—50(—100) cm high. Stem rhizomatous at base,

plants entirely rhizomatous when juvenile with very short internodes, internodes

extending to 5-15 (—30) cm apart when mature, sub-glabrous to hairy, hairs white

Begonia sect. Sphenanthera from Sumatra 247

or red. Stipules persistent, lanceolate, keeled, the keel sometimes with long hairs,

with a fimbriate filiform extension at the tip, 20-30 x 8-15 mm. Leaves: petiole 6-28

cm long, hairy or with scattered hairs 1-2 mm long: lamina ovate, base cordate with

lobes overlapping, 11—23 = 6-16 cm, smaller on juvenile plants, midrib 8.5-16 cm

long, underside with scattered hairs, denser on the veins, upper surface glabrous or

with scattered hairs, hairs white or sometimes reddish on young leaves, venation

palmate-pinnate; margin denticulate. Inflorescences axillary, cymose, bisexual, male

and female flowers open at the same time; bracts lanceolate, c. 15 mm long, smaller

towards the apex of the inflorescence, margin entire or fimbriate: primary peduncle

2-8 cm, secondary 5—15 mm. Male flowers: pedicels c. 10-15 mm long, hairy; tepals

4, white, outer ovate-orbicular, size c. 18 x 18, with red hairs on the outside (always?):

inner tepals elliptic, 15 x 10 mm, glabrous; androecium symmetric, a loose globose

cluster; stamens bright yellow, c. 80 in number, subequal; filaments longer than the

anthers, 2—3 mm long; anthers narrowly elliptic, 1.5—2 mm long, dehiscing through slits

running nearly the entire length of the anther, connective extended. Female flowers:

pedicels 10-15 mm long; tepals 5, white, subequal, elongate-elliptic, outer c. 17 « 12

mm decreasing to 17 x 8 mm for the innermost, outer sometimes with scattered red

hairs; ovary lobed-triangular in cross section, c. 10—15 mm in diameter, either glabrous

or red-hairy, with 3 pronounced fleshy ridges sometimes developed into fairly distinct

triangular equal to subequal wings c. 5mm long, 3-locular, placentae bifid; styles 3,

large, greenish yellow, bifid, stigmatic surface twice spirally and convolutedly twisted,

stigma persistent. Fruit green to reddish, globose-triangular, pendulous, borne in pairs

or fours, 15—20 x 17-20 mm; with three fleshy ridges or stubby wings up to 5-6 mm

wide; glabrous or with scattered hairs; tip bearing remnants of the styles, which wear

away as they become brittle with age. (Fig. 4 & 5)

Material seen. SUMATRA. Aceh: Boer ni Bias, 31 Aug 1934, C.G.G.J.v. Steenis

6207 (BO); ibid, 1934, C_G.G.J.v. Steenis 6237 (BO): Gajolanden, 6 Feb 1904, R.M.

Pringgo Atmodjo 25 (L); Gunung Kemiri, 23 Aug 1971, K. Jwatsuki et al. 1095 (BO);

Gunung Leuser Nature Reserve, Gunung Ketambe, 7 Aug 1972, de Wilde & de Wilde-

Duyfies 14121 (BO); ibid, 13 Aug 1972, de Wilde & de Wilde-Duyfies 14248 (L);

Gunung Leuser Nature Reserve, Gunung Mamas, 10 May 1962, de Wilde & de Wilde-

Duyfies 16757 (BO, L); Kampong Burni Bies, 3 Sep 1971, K. Awatsuki et al. 1562

(BO). North Sumatra: Asahan, Dolok Si Manoek-manoek, 28 Oct 1936, Rahmat si

Boeea 10651 (A); Berastagi Woods, 15 Feb 1921, H.N. Ridley s.n. (K); ibid, 24 May

1921, J.A. Lorzing 8385 (BO): Gunong Batu Lopang, 8 Jul 1972, W..J.O. de Wilde

13497 (BO [2 sheets], L); Gunung Sibayak, Feb 1921, H.N. Ridley s.n. (K); ibid, 12

May 2007, M. Hughes & D. Girmansyah MH1385 (BO, E): ibid, 12 May 2007, M.

Hughes & D. Girmansyah MH1390 (BO, E); Gunung Sinabung, 14 May 2007, M.

Hughes & D. Girmansyah MH1396 (BO, E); Sipirok, Sibual-buali, 19 May 1993,

J.J. Afriastini 2374 (BO). West Sumatra: Bukik Bulek, 19 Apr 2003—20 Apr 2003,

Anda collectors s.n. (ANDA):; Gunung Merapi, 26 Jul 2009, M@. Hughes & A. Taufiq

MH1569 (ANDA, BO, E); ibid, 22 Jun 1953, J.v. Borssum Waalkes 2191 (BO); ibid,

16 Sep 1918, H.A.B. Bunnemeijer 4644 (BO, L); ibid, 20 Jul 2006, D. Girmansyah &

et al. 772 (BO, E): ibid, 18 Feb 1998, W.S. Hoover & J.M. Hunter 877 (BO [2 sheets]):

Mt. Tandikat, 6 Aug 1988, H. Nagamasu 3002 (ANDA, L): Pajakumbuh, Mt. Sago, 19

Oct 1986, Anda collectors 13 (ANDA).

248 Gard. Bull. Singapore 62 (2) 2011

Fig. 4. Begonia scottii habit (main photo) and fruits from Gunung Merapi, West Sumatra (inset

top left) and from Gunung Kemiri, Aceh (inset lower left). Female flower and ovary (inset

right and bottom right) from Gunung Sorik Merapi, North Sumatra. Photo credits: Deden

Girmansyah and Mark Hughes.

Notes. Despite being quite widely collected, this species was only recently described.

Ridley determined some of his collections of this species to his B. trigonocarpa (=B.

multangula), but the two species differ considerably; B. scottii has smaller, ovate (not

broadly lobed) leaves and often comparatively thin-skinned fruit usually borne in

pairs or fours on slender pedicels (not clusters of c. 10 thicker walled berries on stout

pedicels). Three collections from the northern part of Sumatra (van Steenis 6207 and

6237, de Wilde & de Wilde Duyfjes 13531 [BO duplicate]) are more robust than is

typical for B. scottii, and also have more fruit per infructescence. These specimens

possibly represent some introgression with B. multangula. The de Wilde & de Wilde

Duyfjes 13531 duplicates in L show that this collection is mixed (or at least extremely

variable) as these 2 sheets are B. multangula, although the reddish hairs suggest these

also have been introgressed with B. scottii.

Begonia sect. Sphenanthera from Sumatra 249

Fig. 5. Begonia scottii. A. Inflorescence and open male flower. B. Female flower. C. Mature

plant in fruit. Drawn by: Heather Raeburn.

D0) Gard. Bull. Singapore 62 (2) 2011

Ecology and distribution. Endemic to Sumatra. In moss and montane rain forest in

West Sumatra, North Sumatra and Aceh at 900-2600 m, though most frequently

reported from 1400-1700 m (Fig. 2). In primary and secondary forest, on steep banks,

stream sides or at the base of tree trunks or rotting logs. IUCN category Least Concern.

Excluded name

Begonia robusta var. hirsutior (Miq.) Golding & Kareg., Phytologia 54(7): 499 (1984)

—Platycentrum robustum var. hirsutior Miq., Fl. Ned. Ind., Suppl. 1: 322 (1861).

TYPE: Sumatra, Palembajan, J.E. Teijsmann (not located).

Miquel created this taxon for a specimen he originally cited under Diploclinium

areoatum Mig (=Begonia areolata (Miq.) Miq.). It differs from the type variety of

B. robusta Blume in having ‘foliis minoribus, paniculis longe pedunculatis, ovarii

maturescentis alulis subaequalibus’. The type, collected in West Sumatra by Teijsmann

cannot be located despite a thorough search in B, BO, K, L and U, and the description

is not sufficient to identify the taxon with certainty. It seems unlikely to represent

the sole record of the otherwise entirely Javanese B. robusta, but could possibly be

referable to Begonia multangula.

ACKNOWLEDGEMENTS. The authors thank the RBGE Small Projects Fund, the Davis

Expedition Fund and the Percy Sladen Memorial Trust for financial support. This research was

facilitated by the Indonesian Ministry of Research and Technology (RISTEK), the Indonesian

Institute of Sciences (LIPI), Direktorat Jenderal Perlindungan Hutan dan Konservasi Alam

(DITJEN PHKA), Bandan Pengolola Kawasan Ekosystem Leuser (BPKEL), the Scottish

Government’s Rural and Environment Research and Analysis Directorate, Universitas

Sumatera Utara, Universitas Syiah Kuala and National Parks Board Singapore. Nurainas, A.

Anggara, A. Taufig, R. Nelvita, A. Sumadiyaya and S. Rasnovi and many others are thanked for

their assistance in the field. The curators of herbaria A, ANDA, B, BM, BO, E, K, L, MICH, P,

SING and U are thanked for facilitating access to specimens. We are grateful to two reviewers

for their constructive and helpful comments.

References

Doorenbos, J., Sosef, M.S.M. & de Wilde, J.J.F.E. (1998) The sections of Begonia

including descriptions, keys and species lists (Studies in Begoniaceae VI).

Wageningen Agr. Univ. Pap. 98(2): 1-266.

Hughes, M. (2008) An annotated checklist of Southeast Asian Begonia. UK: Royal

Botanic Garden Edinburgh.

Hughes, M., Girmansyah, D., Ardi, W.H. & Nurainas. (2009) Seven new Begonia

species from Sumatra. Gard. Bull. Singapore 61(1): 29-44.

Kiew, R. (2005) Begonias of Peninsular Malaysia. Kota Kinabalu: Natural History

Publications, Borneo.

Tebbitt, M.C. (2003) Taxonomy of Begonia longifolia Blume (Begoniaceae) and

related species. Brittonia 55(1): 19-29.

Begonia sect. Sphenanthera from Sumatra 251

Tebbitt, M.C., Forrest, L.L., Santoriello, A.. Clement, W. & Swensen, S.M. (2006)

Phylogenetic relationships of Asian Begonia, with an emphasis on the evolution

of rain-ballist and animal dispersal mechanisms in sections Platycentrum,

Sphenanthera and Leprosae. Syst. Bot. 31(2): 327-336.

Gardens’ Bulletin Singapore 62 (2): 253-275. 2011 253

Names and new combinations for Peninsular Malaysian

species of Codonoboea Ridl. (Gesneriaceae)

R. Kiew! and C.L. Lim

Forest Research Institute Malaysia, 52109 Kepong, Selangor, Malaysia

‘ruth@ frim.gov.my (corresponding author)

ABSTRACT. Five species are reinstated in Codonoboea Ridl. and new combinations are made

for 74 species that occur in Peninsular Malaysia that were formerly included in Henckelia

Spreng. Codonoboea albina (Ridl.) Kiew is reinstated at specific rank and var. winkleri (Ridl.)

Kiew as a variety of C. malayana (Ridl.) Kiew. Lectotypes are designated where appropriate.

Keywords. Codonoboea, Gesneriaceae, nomenclature, Peninsular Malaysia, synonymy,

taxonomy, typification

Introduction

Codonoboea Ridl. in its current wider sense is the largest genus of dicotyledonous

herbs in Peninsular Malaysia with 79 named species. This compares with other large

herb genera such as the 45 species in Argostemma Wall., Rubiaceae, 45 in Sonerila

Roxb., Melastomataceae (Turner 1997) and 52 species in Begonia L., Begoniaceae

(Kiew 2005). Botanical exploration of poorly known areas continues to discover new

Codonoboea species, such as the two new species from Gunung Stong, Kelantan

(Kiew 2009) and several others that await description. Codonoboea is confined to

primary forest where it is ubiquitous from the lowlands to the highlands occurring on

granite, sandstone and quartz derived soils or rocks but it is noticeably absent from

limestone and aquatic habitats. It is diverse in regard to habit (although there are no

climbing or epiphytic species), leaf and flower morphology. Its centre of diversity is

Peninsular Malaysia, but species are also found in S Thailand, Sumatra, Singapore,

Borneo, Palawan (the Philippines), Sulawesi and New Guinea (Weber & Burtt 1998).

Ridley (1923) in his Flora of the Malay Peninsula first described Codonoboea

as a genus and included three species that in his opinion did not fit into any of the other

genera. However, Codonoboea was ill-defined and not clearly distinct from the closest

genus, Paraboea Ridl. Indeed the two diagnostic characters, namely peduncles adnate

to the petiole (epiphyllous) and corolla lobes very short and tooth-like were present in

only two of the three species. Burtt (1971) later transferred some Paraboea species to

Didymocarpus Wall.

In 1990, Kiew reduced Codonoboea to a section within Didymocarpus and re-

defined it to include four similar species with epiphyllous inflorescences. However,

Didymocarpus itself underwent re-definition (Weber & Burtt 1983, 1998) that resulted

in the exclusion of species with plagiotropic fruits that split on the adaxial side as

opposed to the orthotropic ones of Didymocarpus s. str. that split both abaxially as well

as adaxially. Based on morphological similarity, the excluded Peninsular Malaysian

254 Gard. Bull. Singapore 62 (2) 2011

species together with Loxocarpus R.Br. species were placed in Henckelia Spreng., a

small genus of 15 species from S India and Sri Lanka, a decision that resulted in 180

new combinations (Weber & Burtt 1998, Vitek et al. 2000).

Recent molecular analyses (Moeller et al. 2009) now show conclusively

that the Indian and Sri Lankan Henckelia species are distinct from the Peninsular

Malaysian species and among the Peninsular Malaysian species, Loxocarpus species

are also distinct from both Henckelia and other morphologically similar genera. These

non-Henckelia and non-Loxocarpus species are now accommodated in the genus

Codonoboea that 1s the remaining validly published name for this group of species and

the generic circumscription is enlarged to accommodate them. For the 79 Peninsular

Malaysian species, this necessitates making new combinations for 74 of the species

and these are provided below.

Materials and methods

In working toward the family Gesneriaceae account for the on-going Flora of Peninsular

Malaysia project, the Peninsular Malaysian taxa have been re-examined through

extensive fieldwork, examination of herbarium specimens including types, and the

literature. New combinations in Codonoboea are made for accepted names together

with references (including recent literature), synonyms and types. Lectotypes are

designated where appropriate. Notes are provided where an explanation is needed. The

species are arranged alphabetically. In due course, the new taxa will be published and,

for all species, distribution in Peninsular Malaysia will be mapped and a conservation

status for each taxon will be assessed and a key for identification provided.

In Peninsular Malaysia, Codonoboea belongs to a group of genera that includes

the seven in the key below, which provides the characters that distinguish between

them.

Key to the Codonoboea group of genera in Peninsular Malaysia

Fertile stamens 4.

Capsules very slender, becoming ribbed and scarcely splitting ...... Didissandra*

Capsules thick, smooth and hard, splitting completely on the adaxial side .........

sabdbabagvedustcsaisa diese touacucdvoceedtooss cee secaseraa eee tac Bachan ae ea ee ee Ridleyandra

Fertile stamens 2.

Capsules orthotropic, splitting adaxially and abaxially.

Stigma distinctly bilobed. (Always on limestone.)

Rosette herbs, inflorescences scapiform, calyx tubular ..................

asedaddcdvehlnwlncbauuS das sre cmns custecc solv Sorceasceseeeee eae eae oe nee an Damrongia

Caulescent herbs, inflorescences often epiphyllous, calyx divided

tO DASE rs... sau clitisek eee eee ee Microchirita

Stigma peltate or rounded (not bilobed). (Never on limestone.) ............

sedwasinsuwbodne duiedsuaocstes cnducee eek subohesee seen coset ae ate tee ene eee Didymocarpus

Capsules plagiotropic, splitting only adaxially.

Capsules slender, cylindric, 1—5 cm long .................:c0:e0000 Codonoboea

Capsules short and distinctly thicker at base, 0.4—0.9 cm long ................

Peninsular Malaysian Codonoboea 255

“cigs :- tac heane a ce aa Akane ee Loxocarpus

*in Borneo, some Didissandra species have 2 fertile stamens.

Codonoboea Ridl.

Fl. Malay Pen. 2 (1923) 533. LECTOTYPE: Codonoboea leucocodon (Ridl.) Ridl.

(Ivanina (1967) Gesneriaceae, the Carpological Review). Synonyms: Didymocarpus

pro parte Ridl. ibid 506. — Didymocarpus sect. Codonoboea (Ridl.) Kiew, Blumea 35

(1990) 171.— Paraboea pro parte Ridl. ibid 527. — Henckelia Spreng. pro parte Weber

& Burtt, Beitr. Biol. Pflanzen 70 (1998) 316-325: Weber in Kubitzki, Fam. Gen. Vasc.

Pl. 7 (2004) 146.

Perennial herbs with continuous growth. Stems usually woody, erect and either

caulescent with an unbranched or branched stem to c. 50 cm tall or short and forming

a rosette, or creeping. Leaves usually opposite, sometimes alternate to spiral, sessile or

petiolate. Inflorescences axillary, with | or several per axil, sometimes epiphyllous;

pedunculate pair-flowered cymes with 3-4 flowers or reduced to a single flower or

branched, paniculate and many-flowered; bracteoles usually small, sometimes absent

or large and conspicuous. Flowers 5-merous, with a small calyx divided almost to

base; corolla tubular with 5 lobes, the lower 3 lobes often longer, large and trumpet-

shaped to narrowly tubular to campanulate to short-tubed and flat-faced; tube often

pale purple, cream-coloured or white with concolourous lobes or with lobes deeply

coloured and contrasting, less commonly red or yellow; frequently with a pair of

yellow or orange-brown nectar guides, rarely with additional stripes or spots; stamens

2, filaments long or short with the anthers positioned in the mouth of the corolla, anthers

coherent, connectives often with a tooth-like appendage; nectary forming a deep or

shallow ring, sometimes with a lobed rim or short on the adaxial side or it does not

surround the ovary base and is tongue-like or is lacking; ovary cylindric, tapering into

the style, stigma either peltate or rounded (but never bilobed or lamellate). Capsules

plagiotropic, splitting adaxially, long, cylindric, 1-5 cm long, usually slender, rarely

curved and slightly thicker towards the base.

1. Codonoboea alba (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus albus Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905)

41, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 754, Fl. Malay Pen. 2 (1923)

517. Synonym: Henckelia alba (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

338. TYPE: Curtis 3299, Peninsular Malaysia, Perak, Gunung Bujang Melaka

[Bujong Malacca], Aug 1898 (lectotype SING, here designated, isolectotype K).

Codonoboea alba var. major (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus albus var. major Ridl., J. Straits Branch Roy. Asiat.

Soc. 44 (1905) 41, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. (1908) 754, Fl. Malay

Pen. 2 (1923) 517. TYPE: Wray 3905, Peninsular Malaysia, Perak, Gunung Bubu,

Mar 1890 (holotype SING).

Note: Vitek et al. (2000) listed this taxon as ‘var. uncertain’. Examination of the

type specimen shows that it belongs to C. a/ba but that the differences in habit

256

Gard. Bull. Singapore 62 (2) 2011

and internodes length justify its status as a variety. Wray’s collection number was

erroneously cited as 3209 in Ridley (1908).

Codonoboea albina (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus albinus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 37, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 751, Fl. Malay Pen.

2 (1923) 512; Henderson, Malay. Wild Flowers Dicot. (1959) 346; Stone, Fed.

Mus. J. 26 (1981) 99. TYPE: Wray (King’s Coll. 8070), Peninsular Malaysia,

Perak, Gunung Batu Putih, Aug 1886 (holotype SING).

Note: Weber & Burtt (1998), followed by Vitek et al. (2000), were in error in

considering this species as synonymous with Codonoboea hispida (Ridl.) Kiew

because it is clearly distinct in its longer petiole and larger lamina with a distinctly

serrulate margin, its paniculate inflorescence with about 7 flowers and its smaller

flower with purple spots. It is therefore reinstated here as a distinct species.

Codonoboea albomarginata (Hemsl.) Kiew, comb. nov.

Basionym: Didymocarpus albomarginatus Hemsl., J. Bot. 25 (1887) 204; Ridley,

J. Straits Branch Roy. Asiat. Soc. 44 (1905) 39, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 74 (1908) 752, Fl. Malay Pen. 2 (1923) 515; Henderson, Malay. Wild Flowers

Dicot. (1959) 347. Synonym: Henckelia albomarginata (Hemsl.) A.Weber, Beitr.

Biol. Pflanzen 70 (1998) 338. TYPE: Wray 65, Peninsular Malaysia, Perak,

Waterfall Hill, Jan 1884 (lectotype K, here designated, isotype SING).

Codonoboea alternans (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus alternans Ridl., J. Straits Branch Roy. Asiat. Soc.

44 (1905) 37, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 751, Fl. Malay

Pen. 2 (1923) 513. Synonym: Henckelia alternans (Ridl.) A.Weber, Beitr. Biol.

Pflanzen 70 (1998) 338. TYPE: Curtis 3733, Peninsular Malaysia, Perak, Gunung

Bujang Melaka [Bujong Malacca], Dec 1895 (lectotype SING, here designated,

isolectotype K).

Note: Among the syntypes, this specimen is selected because it 1s deposited in

more than one herbarium.

Codonoboea anthonyi (Kiew) C.L.Lim, comb. nov.

Basionym: Didymocarpus anthonyi Kiew, Gard. Bull. Sing. 44 (1992) 24.

Synonym: Henckelia anthonyi (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

339. TYPE: Kiew RK 2700, Peninsular Malaysia, Terengganu, Ulu Besut

(holotype KEP, isotypes K, SING).

Codonoboea ascendens (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus ascendens Ridl., J. Linn. Soc. 32 (1896) 512, J. Straits

Branch Roy. Asiat. Soc. 44 (1905) 45, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74

(1908) 756, Fl. Malay Pen. 2 (1923) 517. Synonym: Henckelia ascendens (Ridl.)

A.Weber, Beitr. Biol. Pflanzen 70 (1998) 339. TYPE: Haviland s.n., Peninsular

Malaysia, Perak, Tapah [Tapa], 1894 (lectotype SING, here designated;

isolectotype BM).

Peninsular Malaysian Codonoboea DST

10.

iva

13.

Codonoboea atrosanguinea (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus atrosanguineus Ridl., Trans. Linn. Soc, ser. 2, Bot. 3

(1893) 328, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 47, J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist. 74 (1908) 758, Fl. Malay Pen. 2 (1923) 518; Kiew, Gard. Bull.

Sing. 42 (1989) 49. Synonym: Henckelia atrosanguinea (Ridl.) A.Weber, Beitr.

Biol. Pflanzen 70 (1998) 340. TYPE (lecto Kiew, 1989): Ridley s.n., Peninsular

Malaysia, Pahang, Sungai Tahan (lectotype SING).

Codonoboea bombycina (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus bombycinus Ridl., J. Straits Branch Roy. Asiat. Soc.

44 (1905) 48, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 759, Fl. Malay

Pen. 2 (1923) 518. Synonym: Henckelia bombycina (Ridl.) A.Weber, Beitr. Biol.

Pflanzen 70 (1998) 340. TYPE: King’s Coll. 10587, Peninsular Malaysia, Perak,

Ulu Bubong, Jul 1886 (lectotype SING, here designated).

Codonoboea breviflora (Ridl.) Kiew, comb. nov.

Basionym: Didissandra breviflora Ridl., Bull. Misc. Inform. Kew 1926 (1926)

474. Synonyms: Didymocarpus breviflorus (Ridl.) A.Weber & Kiew, Gard. Bull.

Sing. 41 (1988) 7, fig. 4d-f & Sa—b. — Henckelia breviflora (Ridl.) A.Weber, Beitr.

Biol. Pflanzen 70 (1998) 340. TYPE: Hume 8437, Peninsular Malaysia, Selangor,

Ulu Gombak, 23 Sep 1921 (holotype SING).

Codonoboea caelestis Ridl.

Bull. Misc. Inform. Kew 1929 (1929) 259. Synonyms: Didymocarpus caelestis

(Ridl.) Kiew, Blumea 35 (1990) 175. Henckelia caelestis (Ridl.) A.Weber,

Beitr. Biol. Pflanzen 70 (1998) 341. TYPE (lecto Kiew, 1990): Henderson

19683, Peninsular Malaysia, Kelantan, Sungai Keteh at Gua Ninik (lectotype K,

isolectotypes CGE, SING).

Codonoboea calcarea (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus calcareus Ridl., Bull. Misc. Inform. Kew 1929 (1929)

258; Kiew, Malay. Nat. J. 48 (1995) 204. Synonym: Henckelia calcarea (Ridl.)

A.Weber, Beitr. Biol. Pflanzen 70 (1998) 341. TYPE (lecto Kiew, 1995): Nur &

Foxworthy SFN 12192, Peninsular Malaysia, Kelantan, Gunung Stong [Gunong

Sitong], 6 Mar 1924 (lectotype K; isolectotype SING).

Codonoboea castaneifolia (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus castaneifolius Ridl., J. Straits Branch Roy. Asiat. Soc.

86 (1922) 302, Fl. Malay Pen. 2 (1923) 514; Kiew, Malay. Nat. J. 48 (1995)

205. Synonym: Henckelia castaneifolia (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 341. TYPE: Yapp 438, Peninsular Malaysia, Perak, Gunung Inas (lectotype

K, here designated; isolectotype CBE).

Codonoboea codonion (Kiew) C.L.Lim, comb. nov.

Basionym: Didymocarpus codonion Kiew, Gard. Bull. Sing. 42 (1989) 49.

Synonym: Henckelia codonion (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

342. TYPE: Kiew B.H. RK 1204, Peninsular Malaysia, Pahang, Kuala Kenyam,

30 Sep 1982 (holotype KEP, isotype SING).

258

14.

Gard. Bull. Singapore 62 (2) 2011

Codonoboea corneri (Kiew) Kiew, comb. nov.

Basionym: Didymocarpus corneri Kiew, Blumea 35 (1990) 172, figs. 2 & 4.

Synonym: Henckelia corneri (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

342. TYPE: Kiew RK 2655, Peninsular Malaysia, Terengganu, W of Chukai

[Chukei], close to Sungai Nipah [Sg. Nipa], 5 May 1988 (holotype L, isotypes K,

KEP, SING).

Codonoboea craspedodroma (Kiew) Kiew, comb. nov.

Basionym: Didymocarpus craspedodromus Kiew, Malayan Nat. J. 41 (1987)

213. Synonym: Henckelia craspedodroma (Kiew) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 342. TYPE: Kiew RK 1757, Peninsular Malaysia, Johor, Sungai Jasin

(holotype KEP).

Codonoboea crinita (Jack) C.L.Lim, comb. nov.

Basionym: Didymocarpus crinitus Jack, Malayan Misc. 1, 2 (1820) 1, J. Straits

Branch Roy. Asiat. Soc. 44 (1905) 49, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.

74 (1908) 759, Fl. Malay Pen. 2 (1923) 519, fig. 124; Henderson, Malay. Wild

Flowers Dicot. (1959) 348, fig. 325. Synonym: Henckelia crinita (Jack) Spreng.,

Syst. veg., ed. 16; 4, 2 (1827) 13. TYPE: Jack s.n., Peninsular Malaysia, Pulau

Pinang [Penang] (holotype E).

Codonoboea crinita var. elongata (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus crinitus Jack var. elongatus Ridl., J. Straits Branch

Roy. Asiat. Soc. 44 (1905) 50, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908)

760, Fl. Malay Pen. 2 (1923) 520. TYPE: Curtis 3781, Peninsular Malaysia,

Perak, Tapah (specimen not located).

Note: Codonoboea crinita is one of the most variable species in Peninsular

Malaysia and a detailed study is required to understand the variation within this

species and whether sub-specific taxa can be recognised.

Codonoboea crocea (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus croceus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 44, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 756, Fl. Malay Pen. 2

(1923) 516. Synonym: Henckelia crocea (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 343. TYPE: Ridley 9776, Peninsular Malaysia, Perak, Gunung Keledang,

on banks, Aug 1890 (lectotype SING, here designated).

Note: The type 1s selected because it includes both flowering and fruiting material

and is the earlier collection by Ridley himself.

Codonoboea curtisii (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus crinitus var. curtisii Ridl., J. Straits Branch Roy.

Asiat. Soc. 44 (1905) 49, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 759.

Synonyms: Didymocarpus curtisii (Ridl.) Ridl., Fl. Malay Pen. 2 (1923) 520. —

Henckelia curtisii (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 343. TYPE:

Curtis s.n., Peninsular Malaysia, Selangor, above the Gap, Gunung Semangkok

Peninsular Malaysian Codonoboea 259

£9:

20.

21.

ZR.

[Gunong Semangko Pass], May 1902 (lectotype SING, here designated).

Note: Vitek et al. (2000) draw attention to the citation of the basionym by Ridley

(1923) as ‘J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74: 759 (1908)’ as erroneous.

Codonoboea davisonii (Kiew) Kiew, comb. nov.

Basionym: Didymocarpus davisonii Kiew, Malayan Nat. J. 43 (1990) 242.

Synonym: Henckelia davisonii (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

343. TYPE: Kiew RK 2834, Peninsular Malaysia, Pahang, Gunung Lesong, 31

Oct 1989 (holotype KEP, isotype SING).

Codonoboea dawnii (Kiew) Kiew, comb. nov.

Basionym: Didymocarpus dawnii Kiew, Malayan Nat. J. 48 (1995) 201.

Synonym: Henckelia dawnii (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

343. TYPE: Davison D 6, Peninsular Malaysia, Perak, N of E-W Highway, 3 Mar

1995 (holotype KEP).

Codonoboea densifolia (Rid|.) C.L.Lim, comb. nov.

Basionym: Didymocarpus densifolius Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 51, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 761, Fl. Malay Pen.

2 (1923) 521. Synonyms: Paraboea densifolia (Ridl.) M.R.Hend., Gard. Bull.

Straits Settlem. 5 (1930) 79. — Henckelia densifolia (Ridl.) A.Weber, Beitr. Biol.

Pflanzen 70 (1998) 343. TYPE: Lake & Kelsall s.n., Peninsular Malaysia, Johor,

Gunung Janing [Janeng] (holotype SING).

Heterotypic synonyms: Paraboea caerulea Ridl., J. Straits Branch Roy. Asiat.

Soc. 44 (1905) 66, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 772, Fl. Malay

Pen. 2 (1923) 529; non Didymocarpus caeruleus (R.Br.) Koord. — Didvmocarpus

azureus B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 31 (1971) 44. TYPE:

Rostado s.n., Peninsular Malaysia, Terengganu, Bundi, 1904 (holotype SING).

Note: Ridley (1905) cited Kelsall as the collector in the protologue of Didymocarpus

densifolius, but the specimen records Lake & Kelsall as the collectors.

Codonoboea doryphylla (B.L.Burtt) C.L.Lim, comb. nov.

Basionym: Didymocarpus doryphyllus B.L.Burtt, Notes Roy. Bot. Gard.

Edinburgh 23 (1960) 99. Synonyms: Didymocarpus lanceolatus Ridl., J. Fed.

Malay States Mus. 4 (1909) 50, Fl. Malay Pen. 2 (1923) 520, nom. illeg. non

C.B.Clarke (1883). — Henckelia doryphylla (B.L.Burtt) A.Weber, Beitr. Biol.

Pflanzen 70 (1998). TYPE: Robinson & Kloss s.n., Peninsular Malaysia, way to

Gunung Irau (?).

Note: A search in BM, CBE, E, K and SING failed to locate the type specimen,

which was known froma single collection; nor are there other specimens annotated

by Ridley. The status of this species at species level is dubious for Ridley (1923)

himself drew attention to its close similarity to C. fasciata.

260

23.

24.

25.

26.

Gard. Bull. Singapore 62 (2) 2011

Codonoboea ericiflora (Ridl.) Ridl.

Fl. Malay Pen. 2 (1923) 533. Basionym: Didymocarpus ericiflorus Ridl., J. Fed.

Malay States Mus. 6 (1915) 166; Kiew, Gard. Bull. Sing. 42 (1989) 51, Blumea 35

(1990) 173. Synonym: Henckelia ericiflora (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 344. TYPE (lecto Kiew, 1989): Ridley 16283, Peninsular Malaysia,

Pahang, Gunung Tahan, Wray’s camp, Jul 1911 (lectotype K, isolectotype SING).

Codonoboea falcata (Kiew) Kiew, comb. nov.

Basionym: Didymocarpus falcatus Kiew, Malayan Nat. J. 41 (1987) 218, fig. 3.

Synonym: Henckelia falcata (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

344. TYPE: Kiew RK 1726, Peninsular Malaysia, Johor, Gunung Janing Barat

(holotype KEP).

Codonoboea fasciata (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus fasciatus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 50, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 760, Fl. Malay Pen. 2

(1923) 520; Kiew, Gard. Bull. Sing. 42 (1989) 52. Synonym: Henckelia fasciata

(Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 344. TYPE (lecto Kiew, 1989):

Ridley 2169, Peninsular Malaysia, Pahang, Sungai Tahan, 1891 (lectotype K,

isolectotype SING).

Codonoboea flava (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus flavus Ridl., J. Linn. Soc. 32 (1896) 507, J. Straits

Branch Roy. Asiat. Soc. 44 (1905) 34, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74

(1908) 749, Fl. Malay Pen. 2 (1923) 510; Henderson, Malay. Wild Flowers Dicot.

(1959) 346. Synonym: Henckelia flava (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70: 344 (1998). TYPE: Ridley 2914, Peninsular Malaysia, Perak Hills, 1891

(holotype BM).

Heterotypic synonyms: Staurogyne serrulata C.B.Clarke, J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist.74 (1908) 642; Ridley, Fl. Malay Pen. 2 (1923) 563. SYNTYPES:

Wray 1599, Peninsular Malaysia, Perak, Ulu Batang Padang (CAL); Scortechini

s.n., Sine loc. (CAL).

Didymocarpus flavus Ridl. var. purpurascens Ridl., J. Straits Branch Roy. Asiat.

Soc. 44 (1905) 34, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 749, syn. nov.

TYPE: Ridley 11901, Peninsular Malaysia, Perak, Bukit Kapayung, Sungai Siput,

Feb 1904 (lectotype K, here designated, isolectotype SING).

Notes: 1. Staurogyne serrulata was synonymised by Burtt, Notes Roy. Bot. Gard.

Edinburgh 36 (1978) 151. We have not seen the Calcutta specimens so are in no

position to choose a lectotype.

2. Ridley (1905) distinguished Didymocarpus flavus var. purpurescens by its

purplish stem and broad leaves but examination of the type shows that its leaves

are no broader (3.25-4.25 x 2.5—5 cm) than those of the typical variety (7.5—

12.5 x 3.5—5 cm). In species of Codonoboea, the presence or absence of purple

coloration of the stem and leaves often varies between populations of the same

species so is not a good taxonomic character. For these reasons and the fact that it

does not differ in other characters, this variety is reduced to synonymy.

Peninsular Malaysian Codonoboea 261

27.

28.

29.

30.

31.

32.

Codonoboea flavescens (Ridl.) Kiew. comb. nov.

Basionym: Didymocarpus flavescens Ridl., J. Straits Branch Roy. Asiat. Soc.

44 (1905) 35, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 749, Fl. Malay

Pen. 2 (1923) 511. Synonym: Henckelia fiavescens (Ridl.) A.Weber, Beitr. Biol.

Pflanzen 70 (1998) 344. TYPE: Ridley 7585, Peninsular Malaysia, Selangor.

Kuala Kubu (holotype SING).

Codonoboea flavobrunnea (Ridl.) Kiew. comb. nov.

Basionym: Didymocarpus flavobrunneus Ridl., Trans. Linn. Soc. ser. 2, Bot. 3

(1893) 329, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 39, J. Asiat. Soc. Bengal.

Pt. 2, Nat. Hist. 74 (1908) 753, Fl. Malay Pen. 2 (1923) 515: Kiew, Gard. Bull.

Sing. 42 (1989) 52. Synonyms: Henckelia flavobrunnea (Ridl.) A.Weber. Beitr.

Biol. Pflanzen 70 (1998) 344. TYPE (lecto Kiew, 1989): Ridley 2/63. Peninsular

Malaysia, Pahang, Tahan Valley, Jul 1891 (lectotype K. isolectotypes BM. SING).

Heterotypic synonym: Didymocarpus flavobrunneus Ridl. var. montanus Ridl..

J. Fed. Mal. States Mus. 6 (1915) 167, Fl. Malay Pen. 2 (1923) 515: Kiew, Gard.

Bull. Sing. 42 (1989) 52. TYPE (lecto Kiew, 1989): Ridley 16284, Peninsular

Malaysia, Pahang, Gunung Tahan, Wray’s Camp. Jul 1911 (lectotype K.

isolectotype SING).

Codonoboea floribunda (\\.R.Hend.) C.L.Lim, comb. nov.

Basionym: Paraboea floribunda M.R.Hend., Gard. Bull. Sing. 7 (1933) 117.

Synonyms: Didymocarpus floribundus (M.R.Hend.) B.L.Burtt, Notes Roy. Bot.

Gard. Edinburgh 31 (1971) 44. — Henckelia floribunda (M.R.Hend.) A.Weber.

Beitr. Biol. Pflanzen 70 (1998) 345. TYPE: Corner 26022, Peninsular Malaysia.

Terengganu, Kemaman, Bukit Kajang, Sungai Nipah, Jun 1932 (holotype K).

Codonoboea geitleri (A.Weber) C.L.Lim, comb. nov.

Basionym: Didymocarpus geitleri A.Weber, Pl. Syst. Evol. 165 (1989) 95.

Synonym: Henckelia geitleri (A.Weber) A. Weber, Beitr. Biol. Pflanzen 70 (1998)

345. TYPE: Weber & Anthonysamy 860824—3/1, Peninsular Malaysia, Pahang.

Kuantan, Sungai Pandan, 24 Aug 1986 (holotype WU, isotypes WU, KEP).

Codonoboea glabrata (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus glabratus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 38, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 751, Fl. Malay Pen. 2

(1923) 513. Synonym: Henckelia glabrata (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 345. TYPE: King’s Coll. 209, Peninsular Malaysia, Johor, Gunung

Panti, Jun 1880 (lectotype SING, here designated: isolectotype K).

Note: The Singapore sheet is chosen as the lectotype because the K sheet is a

portion of the Singapore one.

Codonoboea grandifolia (Rid\.) Kiew. comb. nov.

Basionym: Paraboea grandifolia (Ridl.) Ridl., Fl. Malay Pen. 2 (1923) 531.

Synonyms: Didymocarpus grandifolius Ridl., J. Linn. Soc. Bot. 38 (1908) 318

non Didymocarpus grandifolius (A.Dietr.) F.G.Dietr. (1834). — Didymocarpus

33:

34.

35:

Gard. Bull. Singapore 62 (2) 2011

tahanicus B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 31 (1971) 46; Kiew,

Malay. Nat. J. 48 (1995) 205, Gard. Bull. Sing. 42 (1989). — Henckelia tahanica

(B.L.Burtt) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 357. TYPE: Wray &

Robinson 5369, Peninsular Malaysia, Pahang, Gunung Tahan (holotype BM fide

Burtt (1971), isotype SING).

Codonoboea heterophylla (Ridl\.) C.L.Lim, comb. nov.

Basionym: Didymocarpus heterophyllus Ridl., Trans. Linn. Soc, ser. 2, Bot. 3

(1893) 329, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 55, J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist. 74 (1908) 763, Fl. Malay Pen. 2 (1923) 522; Kiew, Gard. Bull.

Sing. 42 (1989) 53. Synonym: Henckelia heterophylla (Ridl.) A.Weber, Beitr.

Biol. Pflanzen 70 (1998) 346. TYPE: Ridley 2170, Peninsular Malaysia, Pahang,

Sungai Tahan, 1891 (holotype SING).

Codonoboea hirsuta (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus hirsutus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 48, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 758, Fl. Malay Pen. 2

(1923) 520. Synonym: Henckelia hirsuta (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 346. TYPE: Kings Coll. 715, Peninsular Malaysia, Perak, Gopeng

[Goping], Sep 1880 (lectotype SING, here designated).

Note: King’s Coll. 715 is selected as the lectotype because of confusion with the

numbering of Kings Coll. 2529 in Ridley (1905) and 2829 in Vitek et al. (2000),

which has not been relocated to verify the correct number.

Codonoboea hirta (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus hirtus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 36, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 750, Fl. Malay Pen. 2

(1923) 512. Synonyms: Henckelia hirta (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 346. — Paraboea campanulata Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 65, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 771, Fl. Malay Pen.

2 (1923) 529. — Didymocarpus campanulatus (Rid|.) B.L.Burtt, Notes Roy. Bot.

Gard. Edinburgh 31 (1971) 44, syn. nov. TYPE: Curtis s.n., Peninsular Malaysia,

Selangor, above the Gap, on the road to Simpang Mines, May 1902 (holotype

SING).

Heterotypic synonym: Chirita uniflora Ridl., J. Straits Branch Roy. Asiat.

Soc. 61 (1912) 34, Fl. Malay Pen. 2 (1923) 525. TYPE: Ridley s.n., Peninsular

Malaysia, Selangor, Simpang Mines Track, Apr 1911 (holotype K).

Notes: 1. Ridley (1905) originally described Didymocarpus hirtus from material

without flowers but later (Ridley 1923) described the corolla as ‘nearly | in. long’,

compared with P. campanulata that he described with corollas ‘half an inch long’.

Apart from corolla length, specimens of these two taxa share the same habit, leaf

arrangement and morphology. Weber & Burtt (1998) are correct in suggesting

that ‘this is probably no more than an abnormal specimen of Henckelia with a

very short corolla’. It is here reduced to synonymy.

2. Wood (1974) referred Chirita uniflora to Didymocarpus and Vitek et al. (2000)

Peninsular Malaysian Codonoboea 263

36.

a7.

38.

based on Ridley’s description suggested that it belonged to Henckelia hirta. Based

on examination of the type, Rafidah (2010) confirmed that Chirita uniflora is a

synonym of Codonoboea hirta.

Codonoboea hispida (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus hispidus Ridl., J. Linn. Soc. 32 (1896) 507, J. Asiat.

Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 750, Fl. Malay Pen. 2 (1923) 511;

Henderson, Malay. Wild Flowers Dicot. (1959) 347. Synonyms: Didymocarpus

hispidulus Ridl. (in error), J. Straits Branch Roy. Asiat. Soc. 44 (1905) 35. —

Henckelia hispida (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 346. TYPE:

Curtis 2037, Peninsular Malaysia, Perak, Taiping [Thaiping] Hills, Gunung Hijau,

Sep 1889 (holotype SING).

Heterotypic synonyms: Didymocarpus hispidus Ridl. var. selangorensis Ridl., J.

Straits Branch Roy. Asiat. Soc. 44 (1905) 36 J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.

74 (1908) 750, Fl. Malay Pen. 2 (1923) 512; Stone, Fed. Mus. J. 26 (1981) 99,

syn. nov. TYPE: Curtis 3752, Peninsular Malaysia, Selangor, Gunung Semangkok

[Gunong Semangko], (holotype SING).

Didymocarpus albinellus Ridl., J. Fed. Malay States Mus. 4 (1909) 51, Fl. Malay

Pen. 2 (1923) 512; Henderson, Malay. Wild Flowers Dicot. (1959) 346. TYPE:

Ridley 13679, Peninsular Malaysia, Pahang, Gunung Beremban [Berumban],

Nov 1908 (lectotype SING, here designated, isolectotype K).

Staurogyne macrantha C.B.Clarke, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74

(1908) 642; Ridley, Fl. Malay Pen. 2 (1923) 563. TYPE: King’s Coll. 2417,

Peninsular Malaysia, Perak, Larut (holotype CAL n.v.).

Notes: 1. Didymocarpus hispidus var. selangorensis 1s no more than a white-

flowered form of the typical one that has purple stipes on the lower lobes. It is

therefore here reduced to synonomy.

2. Weber & Burtt (1998) were correct in treating Didymocarpus albinellus as a

synonym. It just represents smaller-leaved plants, perhaps the result of growing

in more exposed conditions.

3. Staurogyne macrantha was synonymised by Burtt, Notes Roy. Bot. Gard.

Edinburgh 36 (1978) 151.

Codonoboea holttumii (M.R.Hend.) C.L.Lim, comb. nov.

Basionym: Paraboea holttumii M.R.Hend., Gard. Bull. Sing. 4 (1927) 54.

Synonyms: Didymocarpus holttumii (M.R.Hend.) B.L.Burtt, Notes Roy. Bot.

Gard. Edinburgh 31 (1971) 44. — Henckelia holttumii (M.R.Hend.) A.Weber,

Beitr. Biol. Pflanzen 70 (1998) 346. TYPE: Holttum 10685, Peninsular Malaysia,

Johor, Gunung Belumut, 25 May 1923 (lectotype SING, here designated).

Note: The SING specimen is selected because Henderson was carrying out

research there. In addition, the specimen at K has not been relocated.

Codonoboea inaequalis (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus inaequalis Ridl., J. Linn. Soc. 32 (1896) 506, J. Straits

Branch Roy. Asiat. Soc. 44 (1905) 41 J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.

264

59:

40.

41.

42.

43.

44,

Gard. Bull. Singapore 62 (2) 2011

74 (1908) 753. Synonym: Henckelia inaequalis (Ridl.) A.Weber, Beitr. Biol.

Pflanzen 70 (1998) 347. TYPE: Curtis 2568, Peninsular Malaysia, Kedah, Pulau

Langkawi, Gunung Machinchang [Chinchang], Sep 1890 (holotype SING).

Codonoboea kelantanensis (Kiew) Kiew, comb. nov.

Basionym: Henckelia kelantanensis Kiew, Gard. Bull. Sing. 61 (2009) 73.

TYPE: Chew et al. FRI 53518, Peninsular Malaysia, Kelantan, Stong Tengah

Forest Reserve, path to Camp Cobra, 8 Feb 2007 (holotype KEP, isotypes E, L).

Codonoboea johorica (Ridl.) Kiew, comb. nov.

Basionym: Didissandra johorica Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 22, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 739, Fl. Malay Pen. 2

(1923) 502. Synonym: Henckelia johorica (Ridl\.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 355. TYPE: Ridley 4175, Peninsular Malaysia, Johor, Gunung Panti

(lectotype SING, here designated; isolectotype K).

Notes: Ridley was carrying out research in Singapore when he described this

species. Therefore, the SING specimen 1s selected as lectotype.

Codonoboea lancifolia (M.R.Hend.) C.L.Lim, comb. nov.

Basionym: Didymocarpus lancifolius M.R.Hend., Gard. Bull. Straits Settlem.

4 (1927) 52. Synonym: Henckelia lancifolia (M.R.Hend.) A.Weber, Beitr. Biol.

Pflanzen 70 (1998) 348. TYPE: Burkill & Haniff 16946, Peninsular Malaysia,

Pahang, gorge of the Sungai Tras near Raub, 12 Nov 1924 (holotype SING).

Codonoboea leiophylla (Kiew) C.L.Lim, comb. nov.

Basionym: Didymocarpus leiophyllus Kiew, Gard. Bull. Sing. 44 (1992) 28.

Synonym: Henckelia leiophylla (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

348. TYPE: Kiew RK 2265, Peninsular Malaysia, Terengganu, Ulu Setiu [Setui]

(holotype KEP, isotype SING).

Codonoboea leucantha (Kiew) Kiew, comb. nov.

Basionym: Didymocarpus leucanthus Kiew, Gard. Bull. Sing. 44 (1992) 31.

Synonym: Henckelia leucantha (Kiew) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 348. TYPE: Kiew RK 2767, Peninsular Malaysia, Selangor, Ulu Ampang

(holotype KEP, isotype SING).

Codonoboea leucocodon (Ridl.) Rid.

Fl. Malay. Pen. 2 (1923) 533. Basionym: Paraboea leucocodon Ridl., J. Fed.

Malay States Mus. 6 (1915) 167. Synonyms: Didymocarpus leucocodon (Ridl.)

Kiew, Gard. Bull. Sing. 42 (1989) 53, Blumea 35 (1990) 175. — Henckelia

leucocodon (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 348. TYPE (lecto

Kiew, 1989): Ridley 16041, Peninsular Malaysia, Pahang, Gunung Tahan, Jul

1911 (lectotype K, isolectotype SING).

. Codonoboea lilacina (Ridl.) Ridl.

Fl. Malay Pen. 2 (1923) 534. Basionym: Didymocarpus lilacinus Ridl., Trans.

Penmsular Malzysian Codonoboea 265

47.

Linn. Soc, ser. 2, Bot. 3 (1893) 330, J. Straits Branch Roy. Asiat. Soc. 44

(1905) 56, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 764: Kiew, Blumea

35 (1990) 174, Gard. Bull. Sing. 42 (1989) 54. Synonym: Henckelia lilacina

(RidL) A. Weber, Beitr. Biol. Pflanzen 70 (1998) 348. TYPE (lecto Kiew, 1989):

Ridley 2165, Penmsular Malaysia, Pahang, Tahan Valley, Jul 1911 (lectotype K.

isolectotype SING).

Codonoboea longipes (C_B_Clarke) Kiew, comb. nov.

Basionym: Didymocarpus longipes C.B.Clarke in A.DC. & C.DC., Monogr.

Phan. 5. 1 (1883) 86; Ridley, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 40.

J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 753, FL Malay Pen. 2 (1923)

521. Synonym: Henckelia longipes (C_B.Clarke) A-Weber, Beitr. Biol. Pflanzen

70 (1998) 349. TYPE: Maingay 2723. Peninsular Malaysia, Johor [Malacca in

error], Gunung Ledang [Mt. Ophir], Aug 1867-68 (lectotype K, here designated,

isolectotype L).

Notes: Among the syntypes. Maingay 2723 1s selected because it 1s a flowering

specimen and the sheet icludes a description of the dissected flower. Early

collectors approached Gunung Ledang from Malacca town but in fact Gunung

Ledang lies within the state of Johor.

Codonoboea malayana ( Hook.f.) Kiew. comb. nov.

Basionym: Didyvmocarpus malayanus Hook.f, Gard. Chron. 20 (1896) 123 & fig.

24. Bot. Mag. (1897) t. 7526; Ridley, J. Straits Branch Roy. Asiat. Soc. 44 (1905)

38, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 752, FL Malay Pen. 2 (1923)

514: Henderson, Malay. Wild Flowers Dicot. (1959) 347, fig. 324. Synonym:

Henckelia malayana (Hook-f.) A-Weber, Beitr. Biol. Pflanzen 70 (1998) 349.

TYPE: Curtis s.n., Peninsular Malaysia, Pulau Pinang [Penang] (holotype K).

Heterotypic synonym: Didymocarpus malayanus Hook.f. var. fasciatus Ridl.. FL

Malay Pen. 2 (1923) 514, syn. nov. TYPE: Robinson s.n., Penmsular Malaysia.

Perak, Gunung Korbu [Kerbau], 16 Mar 1913 (lectotype K. here designated).

Notes: Among the three specimens im the type folder at K. Robinson s.n. is

selected because it is annotated as var. fasciata in Ridley’s hand and its leaves are

conspicuously fasciate even in the dried state. However, whether the leaves have

the broad pale grey band along the midrib varies between and within populations

so cannot be used as a taxonomic character. In addition, the plate of Didymocarpus

malayanus im Botanical Magazine shows the leaves to be clearly fasciate.

Codonoboea malayana is a widespread and variable species so the distinction

between a ‘round-leaved form’ of var. fasciatus versus the typical variety with

leaves “ovate blunt or rounded. ... sometimes lanceolate acute’ does not support

it as a distinct taxon. For these reasons this variety is reduced to synonymy.

Codonoboea malayana (Hookf.) Kiew var. winkleri (Ridl.) Kiew. comb. & stat.

nov.

Basionym: Didymocarpus winkleri Ridl., J. Straits Branch Roy. Asiat. Soc. 50

(1908) 123, Fl. Malay Pen. 2 (1923) 515; Lim et al., Malay Nat. J. 61 (2009)

266

48.

49.

Sir

Gard. Bull. Singapore 62 (2) 2011

178, fig. 3. TYPE: Winkler 1791, Peninsular Malaysia, Negeri Sembilan, Gunung

Angsi, 2 Apr 1908 (lectotype SING, here designated, isolectotype BM).

Notes: Weber & Burtt (1998) followed by Vitek et al. (2000) reduced Didvmocarpus

winkleri to synonomy with D. malayanus without giving any reason for their

decision. Apart from its pure white flowers, var. winkleri is distinct from var.

malayana in its erect habit and floriferous inflorescences with about 10 flowers.

Codonoboea marginata (C.B.Clarke) C.L.Lim, comb. nov.

Basionym: Didymocarpus marginatus C.B.Clarke in A.DC & C. DC, Monogr.

Phan. 5, 1 (1883) 96, Fl. Malay Pen. 2 (1923) 516. Synonym: Henckelia

marginata (C.B.Clarke) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 349. TYPE:

Lobb 184, Peninsular Malaysia, ‘Malacca’ (lectotype ‘Luzon’ K, here designated;

isolectotype ‘Malacca’ K).

Heterotypic synonym: Didymocarpus ophirensis Ridl., J. Straits Branch Roy.

Asiat. Soc. 44 (1905) 43, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 755.

TYPE: Ridley 3185, Peninsular Malaysia, Johor [Malacca] lower part of Gunung

Ledang [Mount Ophir] (holotype SING).

Notes: 1. There are two sheets of Lobb 184, one annotated ‘Malacca’, the other

‘Luzon’, which is crossed out and replaced by ‘Malacca’. No doubt both were

collected in the Malacca area, probably from Gunung Ledang (Mt. Ophir), which

is actually in Johor. This ‘Luzon’specimen is selected as the lectotype because it

has both flowers and fruits (the other specimen has only fruits).

2. It is likely that when Ridley was working on his Flora at Kew, he had the

opportunity to examine the Lobb specimens of Didymocarpus marginatus because

in Ridley (1923) he treated his D. ophirensis as a synonym.

Codonoboea miniata (Kiew) C.L.Lim, comb. nov.

Basionym: Didymocarpus miniatus Kiew, Novon 5 (1995) 40. Synonym:

Henckelia miniatus (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 350. TYPE:

Kiew RK 3792, Peninsular Malaysia, Terengganu, Bukit Bauk (holotype KEP,

isotypes L, SING).

. Codonoboea modesta (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus modestus Ridl., J. Fed. Malay States Mus. 6 (1915)

53, Fl. Malay Pen. 2 (1923) 513. Synonym: Henckelia modesta (Ridl.) A.Weber,

Beitr. Biol. Pflanzen 70 (1998) 350. TYPE: Dyak Coll. s.n., Peninsular Malaysia,

Gunung Korbu [Gunong Kerbau], 1913 (holotype K).

Codonoboea nitida (Kiew & A.Weber) Kiew, comb. nov.

Basionym: Didymocarpus nitidus Kiew & A.Weber, Gard. Bull. Sing. 41 (1988)

4, fig. 3 & 4a-c. Synonym: Henckelia nitida (Kiew & A.Weber) A.Weber,

Beitr. Biol. Pflanzen 70 (1998) 351. TYPE: Weber & Anthonysamy 840711-1/1,

Peninsular Malaysia, Selangor, Gunung Bunga Buah, E of summit, Jul 1984

(holotype WU, isotypes E, K, KEP, KLU, L, WU).

Peninsular Malaysian Codonoboea 267

52.

53.

54.

55.

56.

Codonoboea nivea Kiew

Malayan Nat. J. 41 (1987) 210. Synonyms: Didymocarpus niveus (Kiew) Kiew,

Blumea 35 (1990) 174. — Henckelia nivea (Kiew) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 351. TYPE: Kiew B.H. KBH 86-33, Peninsular Malaysia, Pahang, Ulu

Kinchin, Sungai Damong Kechil (holotype KEP, isotypes K, L, SING).

Codonoboea parviflora (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus parviflorus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 33, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 748, Fl. Malay Pen. 2

(1923) 510. Synonym: Henckelia parvifiora (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 352. TYPE: Ridley 11970, Peninsular Malaysia, Negeri Sembilan,

Gunung Angsi, 1914 (lectotype SING, here designated; isolectotype K).

Codonoboea pauziana (Kiew) Kiew, comb. nov.

Basionym: Henckelia pauziana Kiew, Gard. Bull. Sing. 61, 1 (2009) 74, fig. 1.

TYPE: Chew et al. FRI 53513, Peninsular Malaysia, Kelantan, Stong Tengah

Forest Reserve, Batu Hampar to Cobra Camp, 8 Feb 2007 (holotype KEP [flower],

isotypes KEP [fruits], K, L, SAN, SING).

Codonoboea pectinata (Oliv.) Kiew, comb. nov.

Basionym: Didymocarpus pectinatus C.B.Clarke ex Oliv. in Hook., Icon. Pl.

23 (1892) ¢. 2246; Ridley, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 53, J.

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1909) 762, Fl. Malay Pen. 2 (1923) 521.

Synonym: Henckelia pectinata (Oliv.) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

352. TYPE: Kings Coll. 10711, Peninsular Malaysia, Perak, Aug 1886 (lectotype

K, here designated, isolectotype SING).

Notes: The type collection is recorded from limestone rocks. Unfortunately, the

locality of this Perak plant is not recorded, so it is not possible to confirm whether

this is correct. So far in Peninsular Malaysia no Codonoboea species is recorded

from limestone and in Selangor C. pectinata grows on granite rocks. The Kew

specimen is selected as the lectotype because the sheet includes a description of

the species and a rough flower dissection.

Codonoboea platypus (C.B.Clarke) C.L.Lim, comb. nov.

Basionym: Didymocarpus platypus C.B.Clarke, in A.DC. & C.DC., Monogr.

Phan. 5, | (1883) 94; Ridley, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 46,

J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 757, Fl. Malay Pen. 2 (1923)

517; Henderson, Malay. Wild Flowers Dicot. (1959) 348; Kiew, Malay Nat.

J. 41 (1987) 220. Synonym: Henckelia platypus (C.B.Clarke) A.Weber, Beitr.

Biol. Pflanzen 70 (1998) 352. TYPE: Griffith 3825, Peninsular Malaysia, Melaka

[Malacca] (lectotype K, here designated).

Notes: Griffith 3825 is chosen as the lectotype because among the syntypes it is

the only specimen with both flowers and fruits and which shows both the upper

and lower leaf surfaces.

268

58.

60.

61.

Gard. Bull. Singapore 62 (2) 2011

. Codonoboea polyanthoides (Kiew) C.L.Lim, comb. nov.

Basionym: Didymocarpus polyanthoides Kiew, Gard. Bull. Sing. 42 (1989) 56.

Synonym: Henckelia polyanthoides (Kiew) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 352. TYPE: Kiew RK 2480, Peninsular Malaysia, Pahang, Sungai Teku,

29 Mar 1987 (holotype KEP).

Codonoboea primulina (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus primulinus Ridl., J. Fed. Malay States Mus. 10 (1922)

250, Fl. Malay Pen. 2 (1923) 522; Kiew, Malay. Naturalist 37, 2 (1983) 6, fig.

Synonym: Henckelia primulina (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

352. TYPE: Ridley s.n., Peninsular Malaysia, Selangor, Klang Gates, Bukit

Lompat Bayan, 2 Jan 1921 (lectotype SING, here designated; isotype K).

Codonoboea pulchella (Ridl\.) C.L.Lim, comb. nov.

Basionym: Didymocarpus pulchellus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 44, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 755, Fl. Malay Pen. 2

(1923) 516. Synonym: Henckelia pulchella (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 353. TYPE: Machado H.B.S. 11629, Peninsular Malaysia, Pahang,

Kuala [Kwala] Lipis, May 1913 (lectotype SING, here designated; isotype K).

Codonoboea pumila (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus pumilus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 56, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 763, Fl. Malay Pen.

2 (1923) 523; Henderson, Malay. Wild Flowers Dicot. (1959) 349. Synonym:

Henckelia nana A.Weber, Beitr. Biol. Pflanzen 70 (1998) 350. TYPE: Burn-

Murdoch s.n., Peninsular Malaysia, Pahang, Gunung Semangkok [Semangko

Pass], Feb 1904 (lectotype SING, here designated).

Note: Weber in Weber & Burtt (1998) re-named this species Henckelia nana

because another species was already named H. pumila. Since ‘pumila’ has not

been used in Codonoboea, the original specific name is retained for this species.

Codonoboea puncticulata (Rid\.) C.L.Lim, comb. nov.

Basionym: Didymocarpus puncticulatus Ridl., J. Linn. Soc. 32 (1896) 510, J.

Straits Branch Roy. Asiat. Soc. 44 (1905) 55, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 74 (1908) 763, Fl. Malay Pen. 2 (1923) 522; Kiew, Malay Nat. J. 41 (1987)

220. Synonym: Henckelia puncticulata (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 353. TYPE: Ridley s.n., Peninsular Malaysia, Johor, Gunung Panti, Dec

1892 (holotype SING, photo K).

Heterotypic synonym: Didymocarpus perditus Ridl., J. Straits Branch Roy.

Asiat. Soc. 44 (1905) 54, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 763,

Fl. Malay Pen. 2 (1923) 522. TYPE: Ridley s.n., Singapore, Selitar, 3 Nov 1889

(holotype SING).

Notes: Didymocarpus perditus was described from Singapore where Ridley

(1905) discovered two plants in 1889. However, by 1905 he described the locality

as destroyed by cultivation and thought the species probably extinct. Kiew (1987)

Peninsular Malaysian Codonoboea 269

62.

63.

64.

65.

showed that it is conspecific with Codonoboea puncticulata from Peninsular

Malaysia. It has never been collected again from Singapore so is certainly extinct

there.

Codonoboea pyroliflora (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus pyroliflorus Ridl., Trans. Linn. Soc, sen 2, Bot.

3 (1893) 330; Kiew, Gard. Bull. Sing. 42 (1989) 57. Synonyms: Paraboea

pyrolifiora (Ridl.) Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905) 67, J. Asiat.

Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 772, Fl. Malay Pen. 2 (1923) 529. —

Henckelia pyroliflora (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 353. TYPE

(lecto Kiew 1989): Ridley 2164, Peninsular Malaysia, Pahang, Sungai Tahan, Jul

1891 (lectotype K, isolectotypes BM, SING).

Codonoboea quinquevulnera (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus quinquevulnerus Ridl., Trans. Linn. Soc, ser 2, Bot. 3

(1893) 328, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 47, J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist. 74 (1908) 758, Fl. Malay Pen. 2 (1923) 518; Kiew, Gard. Bull.

Sing. 42 (1989) 58. Synonym: Henckelia quinquevulnera (Ridl.) A.Weber, Beitr.

Biol. Pflanzen 70 (1998) 353. TYPE (lecto Kiew, 1989): Ridley 2753, Peninsular

Malaysia, Pahang, Sungai Tahan, Aug 1891 (lectotype K, isolectotype SING).

Codonoboea ramosa (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus ramosus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 34, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 749, Fl. Malay Pen. 2

(1923) 511. Synonym: Henckelia ramosa (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 353. TYPE: Wray 868, Peninsular Malaysia, Perak, Gunung Batu Putih

(holotype SING).

Codonoboea reptans (Jack) C.L.Lim, comb. nov.

Basionym: Didymocarpus reptans Jack, Malayan Misc. 1, 5 (1820) 3; Ridley, J.

Straits Branch Roy. Asiat. Soc. 44 (1905) 42, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist.

74 (1908) 754, Fl. Malay Pen. 2 (1923) 515; Henderson, Malay. Wild Flowers

Dicot. (1959) 346, fig. 323. Synonym: Henckelia reptans (Jack) Spreng., Syst.

veg. ed. 16; 4, 2 (1827) 14. TYPE: Wallich 1830, Peninsular Malaysia, Pulau

Pinang [Penang], 1830 (neotype K).

Notes: Codonoboea reptans is a very variable species with wide ecological

amplitude from the lowlands to mountains. The neotype is selected because it

was also collected from the type locality in Pulau Pinang.

Only var. monticola is distinct from other subspecific taxa of C. reptans by its

much narrower leaves. Whether var. modesta and var. violascens are distinct from

the typical variety needs more field observations of variation between and within

populations.

Codonoboea reptans var. modesta (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus reptans var. modestus Ridl., Fl. Malay Pen. 2 (1923)

516. Synonym: Didymocarpus modestus Ridl., J. Straits Branch Roy. Asiat. Soc.

66.

67.

Gard. Bull. Singapore 62 (2) 2011

82 (1920) 186, nom. illeg., non D. modestus Redl. (1915). TYPE: Ridley s.n.,

Peninsular Malaysia, Selangor, Ulu Gombak (lectotype K, here designated).

Note: Ridley’s specimen is selected as lectotype because it includes a brief

description and drawing of the flower.

Codonoboea reptans var. monticola (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus reptans var. monticolus Ridl., J. Linn. Soc. 32 (1896)

511, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 42, J. Asiat. Soc. Bengal, Pt.

2, Nat. Hist. 74 (1908) 755, Fl. Malay Pen. 2 (1923) 516. Synonym: Henckelia

reptans var. monticola (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 354.

TYPE: Ridley 2913, Peninsular Malaysia, Perak, Larut Hills, 1892 (lectotype

BM, here designated).

Note: Among the specimens that Ridley cited, this specimen is selected as the

lectotype because Ridley (1905) singled out the Larut Hills population as typical

of this variety.

Codonoboea reptans vat. violascens (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus reptans var. violascens Ridl., J. Straits Branch Roy.

Asiat. Soc. 44 (1905) 43, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 755,

Fl. Malay Pen. 2 (1923) 516. TYPE: Ridley 7583, Peninsular Malaysia, Selangor,

Rawang, May 1891 (lectotype SING, here designated; isotype K).

Codonoboea ridleyana (B.L.Burtt) Kiew, comb. nov.

Basionym: Didymocarpus ridleyanus B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh

23 (1960) 99; Kiew, Gard. Bull. Sing. 42 (1989) 59. Synonyms: Didymocarpus

grandiflorus Ridl., J. Fed. Malay States Mus. 6 (1915) 167, Fl. Malay Pen. 2

(1923) 523, non Didymocarpus grandiflorus (Wall.) A.Dietr. ex Steud. —

Henckelia ridlevana A.Weber, Beitr. Biol. Pflanzen 70 (1998) 354. TYPE: Ridley

s.n., Peninsular Malaysia, Pahang, Gunung Tahan, in forest by the stream below

Wray’s camp, Jul 1911 (holotype K).

Notes. 1. For reasons that are not at all clear, Weber considered Didymocarpus

ridleyanus B.L.Burtt as an illegitimate name and on transferring this species to

Henckelia recorded it as H. ridlevana A.Weber nom. nov. However, Didymocarpus

grandiflorus (Wall.) A.Dietr. ex Steud. is a legitimate name based on Chirita

grandiflora Wall. Correctly, Weber’s combination should have been Henckelia

ridleyana (B.L.Burtt) A.Weber.

2. Although reported to be in K (Kiew 1989, Vitek et al. 2000), no specimens of

this species could be located in the collection in a recent (2010) search. Nor are

there any at SING.

Codonoboea robinsonii (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus robinsonii Ridl., J. Linn. Soc, Bot. 38 (1908) 318,

Fl. Malay Pen. 2 (1923) 513; Kiew, Gard. Bull. Sing. 42 (1989) 59. Synonym:

Henckelia robinsonii (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 354. TYPE

Peninsular Malaysian Codonoboea par fi |

68.

69.

70.

WA.

dz.

(lecto Kiew, 1989): Wray & Robinson 5470, Peninsular Malaysia, Pahang,

Gunung Tahan, 5 Jul 1905 (lectotype K, isolectotypes BM. SING).

Codonoboea rubiginosa (Ridl.) C.L.Lim, comb. nov.

Basionym: Paraboea rubiginosa Ridl., J. Linn. Soc. 38 (1908) 319, FI. Malay

Pen. 2 (1923) 530. Synonyms: Didymocarpus rubiginosus (Ridl.) B.L.Burtt,

Notes Roy. Bot. Gard. Edinburgh 31 (1971) 44; Kiew, Gard. Bull. Sing. 42 (1989)

59. — Henckelia rubiginosa (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 355.

TYPE (lecto Kiew, 1989): Wray & Robinson 5390, Peninsular Malaysia, Pahang,

Gunung Tahan, 3 Jun 1905 (lectotype K, isolectotype SING).

Codonoboea rugosa (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus rugosus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 45, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 756, Fl. Malay Pen. 2

(1923) 517. Synonym: Henckelia rugosa (Ridl.) A.Weber, Beitr. Biol. Pflanzen

70 (1998) 355. TYPE: Gimlette s.n., Peninsular Malaysia, Kelantan, Kuala

[Kwala] Lebir (holotype SING).

Heterotypic synonym: Didymocarpus lithophilus Kiew, (Gard. Bull. Sing. 42

(1989) 54, nomen), Gard. Bull. Sing. 44 (1992) 38. TYPE: Ridley 2/52, Peninsular

Malaysia, Pahang, Sungai Tahan, Aug 1891 (holotype K, isotype SING).

Codonoboea salicina (Rid|.) C.L.Lim, comb. nov.

Basionym: Didymocarpus salicinus Ridl., Trans. Linn. Soc. ser 2, Bot., 3 (1893)

329, J. Straits Branch Roy. Asiat. Soc. 44 (1905) 52, J. Asiat. Soc. Bengal, Pt.

2, Nat. Hist. 74 (1908) 761; Kiew, Gard. Bull. Sing. 42 (1989) 60. Synonyms:

Paraboea salicina (Ridl.) Ridl., Fl. Malay Pen. 2 (1923) 530. — Henckelia salicina

(Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 355. TYPE (lecto Kiew, 1989):

Ridley 2166, Peninsular Malaysia, Pahang, Sungai Tahan, Aug 1891 (lectotype K,

isolectotypes BM, SING).

Heterotypic synonyms: Didymocarpus filicifolius Ridl., J. Fed. Malay States

Mus. 6 (1916) 116, Fl. Malay Pen. 2 (1923) 530; Kiew, Gard. Bull. Sing. 42

(1989) 60. — Paraboea filicifolia (Ridl.) Ridl., Fl. Malay Pen. 2 (1923) 530. TYPE

(lecto Kiew, 1989): Ridley 16059, Peninsular Malaysia, Pahang, Gunung Tahan,

Padang, Jul 1911 (lectotype K, isolectotypes BM, SING).

Codonoboea salicinoides (Kiew) C.L.Lim, comb. nov.

Basionym: Didymocarpus salicinoides Kiew, Gard. Bull. Sing. 44 (1992) 35.

Synonyms: Henckelia salicinoides (Kiew) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 355. — Paraboea salicina (Ridl.) Ridl. var. major Ridl., Fl. Malay Pen. 5

(1925) 325. TYPE (lecto Kiew, 1993): Yapp 193, Peninsular Malaysia, Kelantan,

Kuala Aring (lectotype K, isolectotype CGE).

Codonoboea serratifolia (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus serratifolius Ridl., J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 74 (1909) 761, Fl. Malay Pen. 2 (1923) 521. Synonyms: Didymocarpus

serratus Ridl., J. Straits Branch Roy. Asiat, Soc. 44 (1905) 52, nom. illeg., non

D. serratus R.Br. (1839). — Henckelia serratifolia (Ridl.) A.Weber, Beitr. Biol.

—

iP)

WB:

74.

76.

Til

Gard. Bull. Singapore 62 (2) 2011

Pflanzen 70 (1998) 356. TYPE: Ridley 11922, Peninsular Malaysia, Perak,

Larut Hills, below the Tea Gardens, Feb 1904 (lectotype SING, here designated;

isolectotype K).

Codonoboea soldanella (Ridl.) C.L.Lim, comb. nov.

Basionym: Didymocarpus soldanellus Ridl., J. Straits Branch Roy. Asiat. Soc. 61

(1912) 33, Fl. Malay Pen. 2 (1923) 523. Synonym: Henckelia soldanella (Ridl.)

A.Weber, Beitr. Biol. Pflanzen 70 (1998) 356. TYPE: Haniff s.n., Peninsular

Malaysia, Perak, Gunung Korbu [Kerbau], Jul 1910. cult. B. G. Sing. (holotype

SING).

Codonoboea stolonifera (Kiew) Kiew, comb. nov.

Basionym: Didymocarpus stoloniferus Kiew, Gard. Bull. Sing. 44 (1992) 36.

Synonym: Henckelia stolonifera (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

357. TYPE: Kiew RK 1638, Peninsular Malaysia, Pahang, Gunung Ulu Kali, 25

Mar 1985 (holotype KEP, isotype SING).

. Codonoboea tiumanica (Ridl.) C.L.Lim, comb. nov.

Basionym: Paraboea tiumanica Burkill ex Ridl., Fl. Malay Pen. 2 (1923) 530.

Synonyms: Didymocarpus tiumanicus (Ridl.) B.L.Burtt, Notes Roy. Bot. Gard.

Edinburgh 31 (1971) 44. — Henckelia tiumanica (Burkill ex Ridl.) A.Weber, Beitr.

Biol. Pflanzen 70 (1998) 357. TYPE: Burkill 1142, Peninsular Malaysia, Pulau

Tioman [Tiuman], West of Juara Bay, 29 Jun 1915 (lectotype K, here designated;

isolectotype SING).

Codonoboea urticoides (A.Weber) Kiew, comb. nov.

Basionym: Didymocarpus urticifolius Ridl., Fl. Malay Pen. 2 (1923) 511, non D.

urticifolius (D.Don) Wonisch (1909). Synonym: Henckelia urticoides A.Weber,

Beitr. Biol. Pflanzen 70 (1998) 358. SYNTYPES: Ridley 14277, Peninsular

Malaysia, Perak, Temengor [Temengoh], Jul 1909 (?K); Curtis 1328, Taiping

[Thaiping] Hills, Gunung Hijau, Dec 1887 (?K).

Notes: Its smaller stature and two-flowered inflorescences with larger flowers

place this species in Codonoboea rather than Didymocarpus. Neither Weber

(1998) nor Vitek et al. (2000) typified this species, although Vitek et al. cited

both syntypes as being at K. However, a search there and at SING could not

locate either of the syntypes nor any other specimens annotated as this species.

Therefore it is not possible either to lectotypify or neotypify this species.

Codonoboea venusta (Ridl.) Kiew, comb. nov.

Basionym: Didymocarpus venustus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 51, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 760, Fl. Malay Pen.

2 (1923) 514; Kiew, Malay. Nat. J. 48 (1995) 206. Synonym: Henckelia venusta

(Ridl.) A.Weber, Beitr. Biol. Pflanzen 70 (1998) 358. TYPE (lecto Kiew, 1995):

Curtis 3751, Selangor, Gunung Semangkok, May 1902 (lectotype K, isolectotype

SING).

Heterotypic synonyms: Paraboea pubiflora Ridl., J. Fed. Malay States Mus. 4

Peninsular Malaysian Codonoboea 273

(1909) 51. — Didymocarpus pubifiorus (Ridl.) B.L.Burtt, Notes Roy. Bot. Gard.

Edinburgh 31 (1971) 44. TYPE: Robinson & Kloss s.n., Peninsular Malaysia,

Pahang, Cameron Plateau, Nov 1908 (holotype BM fide Kiew (1995), isotype

SING).

Note: Didymocarpus pubiflorus was reduced to synonomy by Kiew (1995).

78. Codonoboea viscida (Rid\.) Kiew, comb. nov.

Basionym: Didymocarpus viscidus Ridl., J. Straits Branch Roy. Asiat. Soc. 44

(1905) 36, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 750, Fl. Malay Pen. 2

(1923) 512. Synonym: Henckelia viscida (Ridl.) A.Weber, Beitr. Biol. Pflanzen 70

(1998) 359. TYPE: Ridley 9781, Peninsular Malaysia, Perak, Gunung Keledang,

Sep 1898 (holotype SING).

79. Codonoboea yongii (Kiew) C.L.Lim, comb. nov.

Basionym: Didymocarpus yongii Kiew, Gard. Bull. Sing. 42 (1989) 62.

Synonym: Henckelia yongii (Kiew) A.Weber, Beitr. Biol. Pflanzen 70 (1998)

359. TYPE: Kiew RK 2481, Peninsular Malaysia, Pahang, Sungai Teku, 29 Mar

1987 (holotype KEP, isotypes K, L, SING).

Name of uncertain status

Paraboea scortechinii Ridl., J. Straits Branch Roy. Asiat. Soc. 44 (1905) 65, J.

Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74 (1908) 772, Fl. Malay Pen. 2 (1923) 529.

Synonyms: Didymocarpus scortechinii (Ridl.) B.L.Burtt, Notes Roy. Bot. Gard.

Edinburgh 31(1971) 44. — Henckelia scortechinii (Ridl.) A.Weber, Beitr. Biol.

Pflanzen 70 (1998) 356. TYPE: Scortechini s.n., Peninsular Malaysia, Perak (?).

Notes: In the original description, Ridley (1905) noted that the species was

known from ‘a single incomplete specimen’ that had no number or locality. It was

presumably without fruits because no description of them was provided. He noted

that it was allied to P. cordifolia, which unfortunately is an error because there is

no species of this name. Among the species he described under Paraboea, from

the description P. scortechinii is most similar to P. cordata (A.DC.) Ridl. so it is

likely that he intended P. cordata when he wrote P. cordifolia. In 1923, Ridley

again recorded it as ‘very little known’ and no further specimens were cited.

Burtt (1971) transferred all the short-flowered species in Ridley’s Paraboea

sect. Campanulatae that included P. scortechinii to Didymocarpus stating

‘although it has not been possible to confirm the botanical validity of every

species, it seems desirable to provide names in Didymocarpus if they are not

already available’. He did not cite a type so it is not clear whether he had seen any

specimens of this species.

However, since then Didymocarpus has been redefined (Weber & Burtt

1983) to include species with fruits that split along both sutures (among other

characters) while those that split only on the upper side were included in

Henckelia. Didymocarpus cordatus A.DC. remained in Didymocarpus. Weber

274 Gard. Bull. Singapore 62 (2) 2011

& Burtt (1998) transferred D. scortechinii to Henckelia without comment. Vitek

et al. (2000) did not locate the type nor did our search, nor have specimens

subsequently been collected that have been identified as this species.

From the description, P. scortechinii is in fact more similar to D. cordatus in

leaf shape, in its branched panicles and half-inch long flowers, than it is to the

other short-flowered campanulate species transferred first to Didvmocarpus and

then to Henckelia and now to Codonoboea. However, without the fruit the affinity

of this species to either Didymocarpus or Codonoboea cannot be resolved. Its

half-inch long campanulate flower excludes it from belonging to Paraboea as

now redefined by Burtt (1984). Species of Paraboea in Peninsular Malaysia are

restricted to limestone but without locality data the habitat of this species remains

unknown. For all these reasons Paraboea scortechinii 1s listed here as a dubious

name.

ACKNOWLEDGEMENTS. This study was carried out as part of the Flora of Peninsular

-Malaysia Project funded by the Ministry of Science, Technology and Innovation (MOSTI)

through the National Council for Scientific Research and Development (MPKSN) under

Project No. 01-04-01-0000 Khas 2 entitled ‘Safeguarding the Forest Plant Diversity of

Peninsular Malaysia’ and in collaboration with the Royal Botanic Garden Edinburgh. We are

greatly indebted to Michael Moeller for his guidance in the molecular biology analysis and to

D.J. Middleton, A.R. Rafidah and T.L. Yao for discussions on Asian Gesneriaceae, and to L.G.

Saw, R.C.K. Chung and an anonymous reviewer for comments on the manuscript. We are also

grateful to the curators of the BM, CGE, E, K, KLU, L and SING herbaria for permission to

examine their collections.

References

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from south eastern Asia. Notes Roy. Bot. Gard. Edinburgh 31: 35—S2.

Burtt, B.L. (1978) Studies in the Gesneriaceae of the Old World XLIII: Notes on

Malesian Didymocarpus. Notes Roy. Bot. Gard. Edinburgh 36: 151.

Burtt, B.L. (1984) Studies in the Gesneriaceae of the Old World XLVII: Revised generic

concepts for Boea and its allies. Notes Roy. Bot. Gard. Edinburgh 41: 401-452.

Kiew, R. (1987) The herbaceous flora of Ulu Endau, Johore-Pahang, Malaysia,

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201-234.

Kiew, R. (1989) Didymocarpus (Gesneriaceae) on Gunung Tahan, Malaysia. Gard.

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and its species. Blumea 35: 167-176.

Kiew, R. (1995) A new species and section of Didymocarpus (Gesneriaceae) from

Belum and Temengor, Hulu Perak, Peninsular Malaysia. Malayan Nat. J. 48: 201—

207.

Kiew, R. (2005) Begonias of Peninsular Malaysia. Kota Kinabalu, Malaysia: Natural

History Publications, Borneo.

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Kiew, R. (2009) Three new species of Gesneriaceae from Kelantan, Malaysia. Gard.

Bull. Singapore 61: 73-79.

Moeller, M., Pfosser, M., Jang, C.G., Mayer, V., Clark, A., Hollingsworth, M.L., Barfuss,

M.H.J., Wang, Y.Z., Kiehn, M. & Weber, A. (2009) A preliminary phylogeny of the

‘Didymocarpoid Gesneriaceae’ based on three molecular data sets: incongruence

with available tribal classifications. Amer. J. Bot. 96, 5: 989-1010.

Rafidah, A.R. (2010) Revision of Chirita (Gesneriaceae) for Peninsular Malaysia.

Unpublished MSc. Thesis. University Malaya, Kuala Lumpur, Malaysia.

Ridley, H.N. (1905) The Gesneriaceae of the Malay Peninsula. J. Straits Branch Roy.

Asiat. Soc. 44: 1-92.

Ridley, H.N. (1908) Gesneriaceae. In: King & Gamble, Materials for a Flora of the

Malay Peninsula. J. Asiat. Soc. Bengal 74, 2: 729-908.

Ridley, H.N. (1923) Gesneriaceae. F/7. Malay Penin. 2: 495-547.

Turner, I.M. (1997) (“1995”). A catalogue of the vascular plants of Malaya. Gard. Bull.

- Singapore 47: 1—757.

Vitek, E., Weber, A. & Burtt, B.L. (2000) Names, types and current placement of the

species hitherto referred to Didymocarpus, Loxocarpus, Codonoboea, Platyadenia

and Henckelia (Gesneriaceae). Ann. Naturhist. Mus. Wien 102B: 477-530.

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Gardens’ Bulletin Singapore 62 (2): 277-289. 2011 DTG

Studies on Homalomeneae (Araceae) of

Peninsular Malaysia I:

An historical and taxonomic review of

the genus Homalomena (excluding Chamaecladon)

Kiaw Kiaw Ng, P.C. Boyce! and S. Othman

Pusat Pengajian Sains Kajihayat (School of Biological Sciences),

Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Malaysia

‘phymatarum@googlemail.com (corresponding author)

ABSTRACT. An historical and taxonomic review of Homalomena (excluding species assigned

to the Chamaecladon Supergroup sensu Boyce & Wong) for Peninsular Malaysia is presented.

Five species are recognised of which one, H. truncata (Schott) Hook.f. represents a new species

record for the Peninsula. Keys to the Peninsular Malaysian Supergroups and species, and a

taxonomic conspectus, are provided. Three species are illustrated from living plants, and H.

wallichii from the type specimen.

Keywords. Araceae, Homalomena, Supergroup, Peninsular Malaysia

Introduction

Recent publications on Asian Homalomena have focussed on Papuasia (Hay 1999,

Herscovitch & Hay 2003) and East Sunda (Hay & Herscovitch 2002; Boyce & Wong

2008, 2009; Boyce, Wong & Fasihuddin 2010). Until recently, apart from ad hoc

descriptions of new taxa (Baharuddin & Boyce 2005, 2009), little attention has been

paid to the genus in Peninsular Malaysia. Apart from Ridley’s adequate but now out-

dated account (1925), the only complete review is that of Furtado (1939), an account

marred not least by poor and often confused species’ delimitation.

Hay (1999) provided a concise taxonomic and nomenclatural historical review

of Homalomena in Papuasia. More recently, Boyce & Wong (2008) and Ng et al. (in

press) proposed and elaborated informal supergoups and species complexes as a tool

to aid taxonomic study of this speciose and taxonomically ill-understood genus.

This is a precursory account intended to provide a taxonomically and

nomenclaturally stable framework for the Peninsular Malaysian Homalomena species

belonging to the Homalomena and Cyrtocladon Supergroups (sensu Boyce & Wong

2008). From this platform it is intended to tackle the taxonomy of the numerous

undescribed Peninsular Malaysian species. Species assigned to the Chamaecladon

Supergroup (sensu Boyce & Wong 2008), which account for the majority of the

described species in the Peninsula, present considerable taxonomic and nomenclatural

problems and will be the focus of a future study.

278 Gard. Bull. Singapore 62 (2) 2011

History of the genus in Peninsular Malaysia

The first Homalomena species pertinent to Peninsular Malaysia appeared in No.1 of

Jack’s Descriptions of Malayan Plants (Jack 1820) as Calla angustifolia Jack and

C. humilis Jack. Both are now treated as synonyms of Homalomena humilis (Jack)

Hook.f. (Chamaecladon Supergroup — see Boyce & Wong 2008) and fall outside the

scope of this paper.

Griffith (185la) posthumously established new four species, three in

Homalomena, and one in a new genus, Cyrtocladon Griff. (see also Griffith 1851b).

Two of Griffith’s Homalomena species (H. minor Griff. and H. major Griff.), were

based upon incomplete material from the same locality (Pulau Besar, Melaka),

and since publication have been treated as, and here remain, doubtful. Griffith’s

third species, H. rostrata Griff., with which Cyrtocladon sanguinolentum Griff. is

conspecific, is a highly variable colonial helophyte widespread in Sunda and with

an extensive synonymy. It has been treated in an account of helophytic Homalomena

(Wong, Boyce & Fasihuddin 2011).

Schott (1856) recognised nine species for Homalomena, with only those

described by Griffith listed for the Peninsula. Schott retained Griffith’s Cyrtocladon,

but made no mention of the species assigned by Jack to Calla .

Schott (1860) recognized 12 species for Homalomena, and a further nine in

Chamaecladon, which he treated as a genus, and no longer accepted Cyrtocladon.

He listed four Homalomena species for the Peninsula, viz. Griffith’s three species (H.

major, H. minor, H. rostrata), and added H. wallichii Schott, which he had described

from Pinang the previous year (Schott 1859).

Engler (1879) recognized 17 species for Homalomena, but made no increase

to the number, nor altered the status, of species in the Peninsula from that of Schott

(1860). Engler treated Chamaecladon as a genus, with 13 species.

Hooker (1893) increased to 23 the number of species in Homalomena, with

17 recorded for the Peninsula and Singapore, of which 12 are in Chamaecladon,

treated as a section. Of the other five species, Hooker erroneously considered Javan

H. coerulescens Jungh. ex Schott to be present in the Peninsula and within it included

H. major, H, minor, and H. wallichii as synonyms. Notwithstanding that H. minor and

H. major (Griffith 1851) have priority over H. coerulescens (Schott 1856), virtually

all material determined as H. coerulescens by Hooker represents novel taxa. The

remaining species comprise H. paludosa Hook.f., H. rostrata, and H. sagittifolia, now

treated as conspecific (= H. rostrata), and Peninsular Malaysian and southern Thai H.

pontederiifolia Griff. ex Hook.f.

Ridley’s precursory accounts for the Peninsula (Ridley 1902, 1907) listed 21

species with 17, including H. mixta Ridl. (= Furtadoa mixta (Ridl.) M.Hotta — see

Hotta 1985), treated as belonging to sect. Chamaecladon. The taxonomy of Ridley’s

1907 treatment follows that of Hooker (1893) with the exceptions that H. propinqua

Schott was added to the synonymy of H. sagittifolia, and H. pontederiifolia is treated

as a synonym of H. coerulescens. Ridley (1907) made no mention of H. ridleyana

Engl. (Engler 1907), having presumably not yet seen the publication.

Engler (1912) remains the only complete revision of Homalomena to date.

He recognised 80 species, of which 37 are in sect. Chamaecladon. Excluding those

assigned to Chamaecladon, only six species are recorded for Peninsular Malaysia. One

new species, H. curvata Engl., is added but otherwise the account generally follows

Historical and taxonomic review of Homalomena 279

those of Hooker (1893) and Ridley (1907) in listing H. coerulescens (including H.

major, H, minor, and H. wallichii as synonyms), and maintaining H. paludosa, H.

rostrata, and H. sagittifolia as distinct species. Engler resurrected H. pontederiifolia,

and treated H. ridleyana as a synonym of H. paludosa.

Ridley (1925) treated 24 species, with 20 species in sect. Chamaecladon. The

taxonomy is otherwise as that of Engler (1912) with the exception that Ridley reduced

H. curvata to H. coerulescens, and combined H. pontederiifolia as a variety of H.

coerulescens. Homalomena deltoidea Hook.f. (= H. griffithii (Schott) Hook.f.) is

misplaced in sect. Homalomena (‘Eu-Homalomena’ ).

Furtado (1939) recognized 58 species, of which 35 were placed in sect.

Chamaecladon, four in Homalomena (‘Eu-Homalomena’), and 18 in Cyrtocladon,

newly treated as a section. Excluding those assigned to Chamaecladon, Furtado

listed four species for the Peninsula, of which three (H. propinqua, H. rostrata, and

H. sagittifolia) are now treated as a single species: H. rostrata. Furtado attempted

to update Ridley (1925) and Engler (1912), in particular to deal with confusion

surrounding early names published for what is now Indonesia, but in fact his account

created more confusion than it resolved. The biggest problem is Furtado’s hopelessly

muddled treatment of Javan H. rubra Hassk., a species absent from the Peninsula, but

into which Furtado subsumed five Peninsular taxa: H. curvata, H. major, H. minor, H.

pontederiifolia, and H. wallichii.

Conspectus of Homalomena (excluding the Chamaecladon Supergroup)

in Peninsular Malaysia

Homalomena curvata Engl., Pflanzenr., 55(1V.23Da): 53 (1912). TYPE: Malaysia,

Melaka, (“Malakka”’), Wells Hill, bei Kuala Lumpur (but see below), 50 m, Feb 1906,

A. Engler (Reise nach Java und Brit. Indien) n.5292 (holo B!). (Fig. 1)

Notes: Engler’s type locality data are confusing. While Wells Hill (modern Bukit Cina)

is indeed in Melaka, it iies some 120 km SE of Kuala Lumpur. It is probable that

Engler’s labeling led Ridley to not recognise H. curvata as a valid species, since Ridley

appears to have not seen the type and instead based his decision on his collection

(cited by Ridley 1925) of a different species from Weld’s Hill (now renamed Bukit

Nanas, adjacent to Bukit Mahkamah), Kuala Lumpur. It seems that Ridley confounded

Weld’s Hill, Kuala Lumpur with Wells Hill, Melaka. Ironically, Ridley’s Weld’s Hill

collection almost certainly represents an undescribed species. However, the material

is both inadequate and in a poor state of preservation, and thus unsuitable from which

to prepare a description.

A good portion of Bukit Nanas is in the Bukit Nanas Forest Reserve where some

degree of natural vegetation still exists. This is the same hill as St John’s Institution,

Bukit Nanas Convent, and the imposing KL Tower. On the other hand, the present

Bukit Mahkamah used to be called Court Hill (Mahkamah = Court) because the former

High Court Building was situated there (precursor to the Menara Maybank). There

remains a possibility Ridley’s unnamed taxon may have persisted. Bukit Nanas and

Bukit Mahkamah are on opposite sides of a road that used to be called Weld Road

(now Jalan Raja Chulan).

280 Gard. Bull. Singapore 62 (2) 2011

Museum botanicum Berolinense.

A Eoghan, Raine Jere wend Grit. nti eee

sit. Brveteneas curreke Kye

Titus bet deol. f

een?

Fig. 1. Homalomena curvata Engl. A. Plants in habitat. B. Detail of emerging inflorescences.

C. Developing infructescences. D. Detail of abaxial leaf surface showing the diagnostic striate

pellucid secretory canals. E. Holotype (B). A-D from Ng et al. AR-3052. Photo credits: P.C.

Boyce. E from A. Engler 5292. Photo credit: Botanic Garden and Botanical Museum Berlin-

Dahlem, Freie Universitat Berlin. Used with permission.

Historical and taxonomic review of Homalomena 281

An alternative interpretation is that Engler only wrote “Malakka” because there

was no real country called Malaysia then, and he was in fact referring to Weld’s Hill

near Kuala Lumpur (and then misspelling that as ‘Wells Hill’). If that were the case,

the locality formerly called Weld’s Hill (now Bukit Nanas) used to accommodate at

least two Homalomena spp., not impossible by lowland humid forest and Homalomena

standards. Possible corroboration from Melaka and Bukit Nanas material may become

available.

That no one after Engler recognised H. curvataas a distinct species is remarkable.

Critical examination of the holotype (B), and the first author’s recent new collections

from the Jerantut Krau Wildlife Centre, Pahang (Ng AR-3052 & Ng AR-3053) leave us

in no doubt that the conspicuous striate pellucid secretory canals running parallel to

the primary lateral veins on the abaxial side of the leaf blade make H. curvata a species

wholly distinct from any other yet described in Peninsular Malaysia.

Homalomena curvata appears to be an outlying representative of a species

complex otherwise centred on N. Borneo, and from where at least a dozen species, in

the main associated with limestone, await description.

Homalomena pontederiifolia Griff. ex Hook.f., Fl. Brit. Ind. 6: 533 (1893).

LECTOTYPE (selected here): Malaysia, Melaka, Air Panas (‘Ayer Punus’) — see

below, W.Griffith 5964 (lecto K!). (Fig. 2)

Notes: As treated here Homalomena pontederiifolia is the commonest of the larger

species in Peninsular Malaysia, but paradoxically poorly represented in herbaria,

possibly because its large and bulky nature dissuades collection. It is very likely that

more extensive collecting and exhaustive study will reveal that the species comprises

several distinct taxa.

Hooker cites three syntypes, of which Griffith’s collection, here selected as

lectotype, is the only one that completely matches the protologue. Griffith’s locality

(‘Ayer Punus’) is almost certainly the same as modern Taman Rekreasi Air Panas

Jasin, Melaka; the specimen was likely collected not long before Griffith’s death from

a parasitic liver disease in Melaka in February 1845, shortly before his 35" birthday.

Of the other syntypes cited by Hooker, D.F\A. Hervey s.n (K!), although sterile,

represents an undescribed species of the Hanneae Complex (see Ng et al., in press),

based on the marcescent margin of the petiolar sheath, and leaf blades with scattered

glandular punctuations (the latter admittedly only faintly discernible). Fr B. Scortechini

s.n. (CAL! K!) approaches H. pontederiifolia, but the inflorescences are only half

the typical length for the species and the material is otherwise too depauperate to be

unequivocally placed taxonomically.

Homalomena rostrata Griff., Not. Pl. Asiat. 3: 154 (1851) [‘roshalum’]. TYPE:

Malaysia, Melaka, W. Griffith 5989 (holo K!).

Cyrtocladon sanguinolentum Griff., Not. Pl. Asiat. 3: 147 (1851); Chamaecladon

sanguinolentum (Griff.) Schott, Prodr. Syst. Aroid.: 316 (1860). TYPE: Malaysia,

Melaka, W. Griffith 5990 (holo K!).

282

Gard. Bull. Singapore 62 (2) 2011

Fig. 2. Homalomena pontederiifolia Griff. ex Hook.f. A. Plant in cultivation, ex Johor, Malaysia.

B. Inflorescence at female anthesis. C. Inflorescence at female anthesis, spathe artificially

removed. Scale bar = 2

cm. D. Detail of staminate flower zone. Note the well-defined flowers

each with 4-6 anthers and large synconnectives. E. Detail of pistillate flower zone and lower

portion of staminate zone. The lowermost staminate flowers are sterile (staminodes). Note that

interpistillar staminodes equal the pistils in length. A-E from Boyce AR-2355. Photo credits:

P.¢ Boyce.

Historical and taxonomic review of Homalomena Des

Homalomena sagittifolia Jungh. ex Schott, Prodr. Syst. Aroid.: 311 (1860). TYPE:

Malaysia, Sabah, Labuan, Motley s.n. (holo K!); epitype (see Wong et al., 2011):

Indonesia, Java, FF. W. Junghuhn s.n. Wt; Schott Ic., no. 2111 (W!). — Fiche no. 44:

a4 in the microfiche edition.

Homalomena miqueliana Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 126 (1863).

TYPE: Indonesia: Borneo, Kalimantan, P W. Korthals s.n. (holo L!).

Homalomena propinqua Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 280 (1864).

TYPE: Indonesia, Kalimantan, Kalimantan Selatan, Mt. Gintang, PW. Korthals s.n.

(holo L!).

Homalomena beccariana Engl., Bull. Soc. Tose. Ortic. 4: 296 (1879). TYPE: Malaysia,

Sarawak, Kuching, Jul 1865, O. Beccari PB. 260 (holo FI-B!).

Homalomena paludosa Hook.f., Fl. Brit. India 6: 531 (1893). LECTOTYPE: Malaysia,

Perak, Larut, H.H. Kunstler (‘Dr King’s Collector’) 3082 (lecto K! isolecto CAL,

SING!).

Homalomena sagittifolia var. pontederiifolia Ridl., J. Straits Branch Roy. Asiat. Soc.

44: 172 (1905). — Homalomena ridleyana Engl., Bot. Jahrb. Syst. 37: 123 (1907).

TYPE: Malaysia, Borneo, Sarawak, Kuching, E. Bartlett & G.D. Haviland 3134 (holo

SING!).

Homalomena teysmannii Engl., Pflanzenr., 1V, 23Da: 68 (1912). TYPE: Indonesia,

Sumatera, Bangka Island, JE. Teijsmann 3227 (holo B! iso BO!).

Homalomena raapii Engl. Pflanzenr., 1V, 23Da: 73 (1912). LECTOTYPE (see Wong

et al., 2011): Indonesia, Sumatera, H. Raap 235 (lecto BO!).

Homalomena triangularis Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 181 (1922).

TYPE: Indonesia, Sumatera, Deli, 27 Jul 1915, /.4. Lérzing 4071 (holo BO!).

Homalomena sagittifolia var. sumatrana Alderw., Bull. Jard. Bot. Buitenzorg, III, 4:

192 (1922). LECTOTYPE (see Wong et al., 2011): Indonesia, Sumatera, Ophir, Taloe,

10 Apr 1917, H.A.B. Biinnemeijer 129 (lecto BO!).

Homalomena ensiformis Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 335 (1922).

TYPE: Indonesia, Kalimantan, /.G. Hallier 1465 (holo BO! iso L!).

Homalomena miqueliana var. truella Alderw., Bull. Jard. Bot. Buitenzorg, II, 4: 336

(1922). TYPE: Indonesia, Sumatera, Riau Archipelago, Ampoelai, Pulau Bintang, 14

Jun 1919, H.A.B. Biinnemeijer 6200 (holo BO! iso L!).

Homalomena sagittifolia var. angustifolia Furtado, Gard. Bull. Straits Settlem. 10:

228 (1939). LECTOTYPE (see Wong et al., 2011): Malaysia, Johore, Mt Austin, H.N.

Ridley 12018 (\ecto SING! isolecto K!).

284 Gard. Bull. Singapore 62 (2) 2011

Notes: See Wong et al. (2011) for a detailed discussion and illustrations of this species.

Homalomena truncata (Schott) Hook.f., Fl. Brit. India 6: 535 (1893). —Chamaecladon

truncatum Schott, Bonplandia (Hannover) 6: (1858). LECTOTYPE selected here:

Myanmar, Mergui, W. Griffith s.n. (lecto K!). (Fig. 3)

Notes: Schott (1858) cited no material for Chamaecladon truncatum, although the

protologue is explicitly prepared from preserved rather than living plants. Schott’s

description in the Prodromus (Schott 1860) is largely identical to that in Bonplandia,

and further cites two specimens, from Mergui and Tenasserim, both extant in Kew (K).

That chosen as lectotype is autographed by Schott, and is also the better preserved of

the two.

Schott’s placement of H. truncatum in Chamaecladon 1s at odds with the

morphology of the interpistillar staminodes (exceeding the associated pistils), and the

staminate flowers comprising four stamens and synconnectives. It seems likely that

Schott regarded the rather small, unconstricted spathes as more pertinent to placement

than the finer floral details.

This represents a new record for the Peninsula. Homalomena truncata is a

common species in the north of Peninsular Malaysia occurring in subhumid to humid

forest, often, but not exclusively, associated with limestone.

Homalomena wallichii Schott, Bonplandia (Hannover) 7(3): 30 (1859). TYPE:

Malaysia, Pulau Pinang, Aug 1822, N. Wallich EIC 8951 (holo K-W!). (Fig. 4)

Notes: Much as with H. curvata, H. wallichii was reduced to synonymy virtually the

moment it was published, and as with H. curvata one can only speculate as to how

such a distinctive species could have been so treated. The habit of the sterile plant —

with long-petiolate pendent leaf blades (leaf tip directed downwards) recalls that of an

Alocasia — while fertile plants are unmistakable by the exceptionally long peduncle,

with the inflorescence often equalling the preceding petiole. Homalomena wallichii 1s

very uncommon in the wild, and furthermore appears to be restricted to a very small

area of Pinang, fortunately within the Taman Negara Pulau Pinang (“Muka Head N.P.”)

Inadequately known species

Homalomena major Griff. (‘majus’), Notul. Pl. Asiat. 3: 153 (1851). TYPE: Malaysia,

Melaka, Pulau Besar [‘Pulo Bissar in Sylvis collinis’], Sep 1842. W. Griffith 6011 (K).

Homalomena minor Griff. ( minus’), Notul. Pl. Asiat. 3: 152 (1851). TYPE: Malaysia,

Melaka, Pulau Besar [*Pulo Bissar in Sylvis collinis’], Sep 1842. W. Griffith 5961 (K).

Notes: These collections are almost certainly the same species, but the material is

inadequate, seemingly preserved late in anthesis (thus many important floral characters

probably already lost) and now too fragile to enable dissection to assign them to any

known plant. A further problem is that Pulau Besar is now much degraded, although

286 Gard. Bull. Singapore 62 (2) 2011

Fig. 4. Homalomena wallichii Schott. Holotype, K-W. Photo credit: The Trustees, Royal

Botanic Gardens, Kew. Used with permission.

Historical and taxonomic review of Homalomena DReh

certainly an attempt to re-collect at the site must be attempted before these names are

relegated to “nomina obscura”.

Key 1. Peninsular Malaysian Homalomena Supergroups

la. Spathe at most 1.5cm long, often much less, without a constriction. Staminate

flowers each comprising 2—3 stamens not united by a common connective.

Interpistillar staminodes shorter than the pistils. (Mostly small plants and often

ELST TIDES) | co ace ee eee Oe Chamaecladon Supergroup

1b. Spathe more than 2.5 cm long, often much longer, with or without a constriction.

Staminate flowers each comprising 3-4 (rarely 5—6, very rarely 1) stamens united

by a conspicuous common connective. Interpistillar staminodes equaling or

slightly exceeding the pistils. (Mostly medium to large plants and in the Peninsula

ear TMM LICS OIF YEN SLE IKG pee pete te hae sO oe. oo Se Sado ues nied nace odd dc hush seavteasbivcscdadesdeds Ds

2a. Spathe not divided into a lower and upper portion by a constriction. Inflorescence

movement during anthesis comprising simple gaping and closing of the spathe

mo withiviniiially-Mo SpadixX MOVEMENE ...:-....:....200.c<..cccssseececssedersvenccevecveseccncsesess

~ a Zesiglhae 328 cd ge epee EEE OE Homalomena Supergroup (see Key 2)

2b. Spathe divided by a moderate to pronounced constriction into a well-defined

upper (limb) and a lower portion. Inflorescences during anthesis with complex

spathe and spadix movements and often spadix elongation .............::ccceeeeeeeeeeteees

_ os sibughtcce: Sekek oa ie OE Cyrtocladon Supergroup (see Key 3)

Key 2. Peninsular Malaysian Homalomena

la. Flowering plants large, often exceeding | m tall. Spathe 8-14 cm long; spadix 7-13

SEIN ROI ee eee rae LCR LSS. Sects 1c cada dcssoteisnssdiescastuscevessses H. pontederiifolia

lb. Flowering plants of medium size, seldom exceeding 40 cm tall. Spathe 4-6 cm

MMR e SES AUN De CoN MINE nae. 8o ocudes <2. Je chvvactaccego-tcuctec staceeadebectceveeeveressoceedavesverese 2

2a. Leaf blade abaxially with conspicuous striate pellucid secretory canals running

parallel to the primary lateral veins. (S Peninsula as far north as S Pahang.) .......

MEER EE oy 8 Potion ns Sorc des asst seeeceteoueecerdeesessocdsboselascacaseges H. curvata

2b. Leafblade abaxially without striate pellucid secretory canals. (Kedah, Perlis.) .......

ce peutte245089/ 0000300 Ge eee ee H. truncata

Key 3. Peninsular Malaysian Cyrtocladon

la. Colonial helophytes, almost always in peatswamp or the wet facies of kerangas

(kerapah) forest and always in inundated situations ..............:::0:ce H. rostrata

1b. Solitary to clumping forest mesophytes in moist well-drained situations in lowland

Pern mixed dipterocarp LOTESE ...2.....-...0.-..0c.soreseoeenoetees (numerous sp. nov.)

288 Gard. Bull. Singapore 62 (2) 2011

ACKNOWLEDGEMENTS. The first author’s research and fieldwork in Peninsular Malaysia

was supported by a Research University Grant 1001/PBIOLOGI/811132 from the Universiti

Sains Malaysia (USM), Penang, Malaysia. The first author is also grateful for financial support

supplied by a USM Fellowship. The second author wishes to extend thanks to the Keeper and

staff of the Herbarium, Royal Botanic Gardens, Kew, for graciously allowing access to their

collections and for permission to reproduce in this paper images of pertinent specimens. We

wish to express our gratitude to Dr Wong Khoon Meng (SING) for very useful information

regarding Weld’s Hill, and the implications for yet further novel taxa, reproduced above almost

verbatim.

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Ha Noi, Viet Nam

nguyenvandu@ fpt.vn (corresponding author)

> Pusat Pengajian Sains Kajihayat (School of Biological Sciences), Universiti Sains Malaysia,

11800 USM, Pulau Pinang, Malaysia

phymatarum@googlemail.com

ABSTRACT. Pycnospatha arietina Gagnep. (Araceae—Lasioideae) is recorded as a new

species and genus record for Cambodia and Vietnam. An updated species description and a key

to both species of Pycnospatha are provided. The Cambodian plant is figured.

Keywords. Araceae, Cambodia, Laos, Lasioideae, Pycnospatha, Pycnospatha arietina, Vietnam

Introduction

The first author has for over 15 years been undertaking Araceae-related fieldwork in

Vietnam and following easing of access, more recently in Cambodia and Lao. In 2007,

during fieldwork in the Peah Ream National Park, Sihanouk Ville, Cambodia, material

was collected of Pycnospatha arietina Gagnep. (Araceae - Lasioideae), representing a

new genus and species record for Cambodia. This new Cambodian collection locality

is only approximately 30 km N of Phu Quoc island (Vietnam) from where H6 (1993,

2000) illustrated, as Amorphophallus sp., what is undoubtedly also P. arietina. Thus

the genus Pycnospatha, represented by P. arietina, is also present in Vietnam.

Pycnospatha comprises two species. The type, P. palmata Thorel ex Gagnep.

(Gagnepain 1941) was described from Lao P.D.R. based ona collection made by Clovis

Thorel during his 1886-1868 Mekong Delta expedition. Although as yet not recollected

from Lao, P. palmata has more recently been found in NE Thailand (Bogner 2009).

Pycnospatha arietina (Gagnepain 1941, 1942) was described from a collection

of Put Phraisurind, the Thai assistant of A.F.G. Kerr, originating from central Thailand,

but is now known to be widespread but scattered, and never common, in Central &

SE Thailand (see Boyce 1993). Hu (1968) described a third species, P. soerensenii

S.Y.Hu, differing from P. arietina by a less divided leaf blade, an aculeate petiole

epidermis, and by having some flowers with a rudimentary ‘perianth’. In his revision

Bogner (1973) regarded P. soerensenii as a synonym of P. arietina, a stand maintained

here.

292 Gard. Bulli. Singapore 62 (2) 2011

Key to species

la. Tuber to over 10 cm diam. Leaf to 2.5 m tall; mature leaf blade ‘dracontioid’ (see

below), up to 60 x 100 cm. Spathe 9-20 = 3—5 cm; spadix 4-6 = 1—1.5 cm: style

to 6 mm. Fruiis c.2 cm. diam. 2 eee Pycnospatha arietina

lb. Tuber c. 2.5 cm diam. Leaf to 50 cm tall; mature leaf blade palmately trilobed,

up to 35 x 16 cm. Spathe 3—7 = 1.5—3 cm; spadix 2-3 x c. 0.5 cm; style to 2 mm.

Fruits 7-8 mm diam ...5..<...: ee Pycnospatha palmata

Pycnospatha arietina Gagnep., Bull. Soc. Bot. France 88: 512 (1941) & inH. Lecomte,

Fl. Gén. Indo-Chine 6(9): 1110 (1942); Bogner, Oesterr. Bot. Z. 122: 207 (1973):

Boyce in Curtis’s Bot. Mag. 10(3): 121 — 124, Pl. 226 (1993); H6, Illustr. Fl. Vietnam.

3(2): 448, fig. 8337 (as ‘Amorphophallus sp.’) (1993): Mayo, Bogner & Boyce, Gen.

Araceae. 135, Pl. 24, 111C (1997); H6, Illustr. Fl. Vietnam. 3: 363, fig. 9165 (as

- ‘Amorphophallus sp.”) (2000). TYPE: Thailand [‘Siam’], SE77. Sa Kaeo: Watthana

Nakhon [*Watana’], 15 Oct 1928, Put Phraisurind 1949 (holo K 000291745! iso BK!

BKF! K 000291746! K 000291744!). (Fig. 1)

Pycnospatha soerensenii S.Y. Hu, Dansk Bot. Ark. 23: 418, t. 3, f. 1-3 (1968). TYPE:

Thailand, SE61. Chantaburi: Makarn, 8 Apr 1959, T. Sorensen, K. Larsen & B. Hansen

7217 (holo C!).

Geophytic, seasonally dormant, tuberous, moderately robust herb with leaves 80

cm-—2.5 m tall. Tuber semi-globose, 13—17 cm in diam., upper surface somewhat

irregularly flattened with many roots and numerous cylindrical or globose secondary

tubers. Roots stout, 34 mm in diam. Adult leaf usually solitary, rarely two together;

petiole 80-200 cm long, c. 2.5 cm diam. at base, subtended by a triangular cataphyll:

cataphyll 4-13 = 1-4.5 cm, pale green to whitish, spotted reddish brown; greyish green

to white-grey with blackish green to reddish or dark brown mottling and with scattered

to rather dense prickles: petiolar sheath short, c. 5—7 cm long, closed: adult leaf blade

dracontioid (elaborated forms of sagittate, hastate or trisect leaves with the anterior and

posterior divisions highly dissected and subdivided, with the terminal lobes each with

two or more tips; leaf maturation acropetal with the leaf blade exposed from the bud

at a very early stage in its expansion), up to 1 m wide, anterior lobe trilobed with the

central lobe again bilobed, posterior lobes bilobed with secondary and tertiary lobes

irregularly pinnatifid, each terminal lobe with two or sometimes three tips, blade dark

green, upper surface glossy often with a slight bluish green sheen. Peduncle 1.5—3 cm

long, quite stout, c. 13 mm in diam., and oblong in cross-section at base, ivory at base,

red-brown above, enveloped by up to 7 cataphylls; cataphyll triangular to lanceolate,

shortest one 2.5 cm long, 4 cm wide at base, longest one up to 9 cm, and 3.5 cm wide,

white at base, upper part dull red-brown outside, pink-red inside, darker at margin.

Spathe c. 15 cm long in Cambodian material (overall range 9-20 cm long), lower

part convoluted into a tube, c. 10 cm long, c. 4.5 cm diam., margins not overlapping,

opened, reddish to dark brown, light brown and green yellow at base, with dull stripes,

c. 1 cm distant; spathe limb fornicate, 4.5 cm long, 9 cm wide at apex, 5.5 cm wide at

base, apex truncated, acute at tip, dark red-brown with white spots outside, dark brown

Pycnospatha new for Cambodia and Vietnam

inside. Spadix sessile, conical, c. 4.5 cm long, 2.3 cm diam. at base, covered densely

by flowers and fertile to tip. Flowers bisexual, naked, very densely arranged. Stamens

6, 2-3.5 mm long; filaments flat, c. 1 mm long, whitish; anther truncate apically, 2—2.5

mm long, yellow, dehiscing by a longitudinal slit. Gynoecium ovate, 5—7 * c. 0.7 mm,

Fig. 1. Pycnospatha arietina Gagnep. A. Whole plant (the first author holding the plant with

Mr Rattana, Cambodian guide); B. Leaf blade, top view; C. Tuber with secondary tubers; D.

Inflorescence; E. Spadix; F. Infructescences; G. Fruits, cut to reveal seeds. Photo credits: V.D.

Nguyen.

294 Gard. Bull. Singapore 62 (2) 2011

pale grey; style extending far beyond stamens, conical, c. 6 mm long, 1.8 mm wide;

stigma discoid, flat. Ovules 2, anatropous. Infructescence globose, sometimes short

cylindrical, composed of c. 40 berries; fruits ovate, c. 1 cm diam., with persistent

style at the acuminate apex, spiny, violet-brown outside, white inside, 1 seeded. Seeds

kidney-shaped, c. 4 mm diam., testa hard, thick, verrucose, brown.

Distribution: C, E & SE Thailand, SW Vietnam, S Cambodia.

Habitat: Sandy soil in open places in dry semi-evergreen tropical forest, elsewhere

(Thailand) in seasonally dry bamboo forest. 90-450 m asl.

Materials studied: CAMBODIA: Krong Preah Sihanouk, Peah Ream National Park

10°33°45.6”N 103°39714.5”E, 20 Nov 2007, Nguyen Van Du et al., CB-VN 103 (HN!

K!).

THAILAND. Saraburi: Muak Lek, 3 Sep 1928, Put Phraisurind 1857 (BK! K!);

ex Muak Lek, cult. in Bangkok, 3 Nov 1929, 4.F.G. Kerr 17580 (BK! K!). Rayong:

-Khao Chamao N.P., 9 Sep 1994, PC. Boyce 909 (BKF! K barcode no. 60524!).

Chantaburi: Makam, Subvillage No. XIII, 20 Mar 1971, J. Bogner 395 (B! K barcode

nos. 29047.184! 34425! 34429! M! US!); Makam, 8. Apr 1959, 7: Sorensen, K. Larsen

& B. Hansen 7217 (C!). Sa Kaeo: ex Watthana Nakhon cult. in Bangkok, 15 Jan

1930, A.FG. Kerr 18098 (BK! K!); ex Watthana Nakhon cult. in Bangkok, 27 Oct

1930, A.F.;G. Kerr 19875 (BK! K!); Watthana Nakhon, 14 Oct 1928, Put Phraisurind

1908 (BK! K!); Watthana Nakhon, 15 Oct 1928, Put Phraisurind 1949 (BK! BKF! K

barcode nos. 000291745! K 000291746! K 000291744!).

Conservation: The plant is not common in Cambodia and its continued survival

much depends on local government plans if the place, where the plant occurs is left

to unregulated tourist-related or construction programs. Based on IUCN criteria, P.

arietina is Vulnerable (VU) in Cambodia.

Cultivation: In Hanoi P. arietina of Cambodian origin grows well in sandy soil in an

open position. The tubers need to should be taken up and stored during (Vietnam)

winter to prevent them rotting in the cool, wet weather.

Notes: Neither Bogner (1973) nor Boyce (1993) mentioned the presence of secondary

tubers on the surface of the mother tubers, nor the often globose infructescence.

ACKNOWLEDGEMENTS. The first author would like to thank the National Geographic

Society for grant No.8283-07, Mr Tan Jiew Hoe (Singapore) for supporting fieldwork in

Cambodia in 2007, and the Vietnam National Foundation for Science and Technology

Development (NAFOSTED) for grant no. 106.11.38.09.

References

Bogner, J. (1973) Die Gattung Pycnospatha Thorel ex Gagnep. (Araceae). Oesterr.

Bot. Z. 122: 199-216.

Pycnospatha new for Cambodia and Vietnam 295

Bogner, J. (2009) Pycnospatha palmata Thorel ex Gagnep. (Araceae) rediscovered.

Aroideana 32: 2-7.

Boyce, P.C. (1993) Pycnospatha arietina. Curtis's Bot. Mag. 10(3): 121-124, Pl. 226.

Gagnepain, F. (1941) Pycnospatha Thorel mss., nov. gen. Aracearum. Bull. Soc. Bot.

France 88: 511-512.

Gagnepain, F. (1942) Aracées, in: Lecomte, H. (ed) Floré Général de 1’Indo-chine

6(9): 1075-1196. Paris: Masson et Cie.

Ho, P.H. (1993) Araceae. Cay co Viétnam [An Illustrated Flora of Vietnam] 3: 417—

453. Montréal.

Ho, P.H. (2000) Araceae. Cay co Viétnam [An Illustrated Flora of Vietnam] 3: 334—

367. H6 Chi Minh (Saigon).

Hu, S.Y. (1968) Studies in the Flora of Thailand 41-48: Araceae. Dansk Bot. Ark. 23:

409-457.

Gardens’ Bulletin Singapore 62 (2): 297-300. 2011 297

Additional notes on Maclurochloa montana

(Poaceae: Bambusoideae) in Thailand

C. Rattamanee’, S. Sungkaew~>* and Y. Paisooksantivatana'

"Department of Horticulture, Faculty of Agriculture, Kasetsart University, Bangkok, Thailand

* Department of Forest Biology. Faculty of Forestry, Kasetsart University. Bangkok, Thailand

> Center for Advanced Studies in Tropical Natural Resources, Kasetsart University,

Bangkok, Thailand

* fforsws@ku.ac.th (corresponding author)

ABSTRACT. The bamboo genus, Maclurochloa K.M.Wong. hitherto known by a single species

from Peninsular Malaysia, M. montana (Ridl.) K.M.Wong, is here recorded with certainty for

Thailand. Additional details of morphology based on Thai specimens and illustrations are

provided.

Keywords. Bamboos. Maclurochioa, Thailand

Introduction

Maclurochloa is a genus of bamboo (Poaceae: Bambusoideae: Bambusinae) first

established by Wong (1993) and represented by a single species, VM. montana (Ridl.)

K.M.Wong. It is one of several climbing genera recently shown to be distinct from

Bambusa Schreb. (Wong 1993, 1995, 2005: Goh et al. 2010). Maclurochloa montana

was previously regarded as endemic to Peninsular Malaysia and recorded in the states

of Penang, Kedah, Pahang and Selangor, in lower montane forest at 780—1350 m above

sea level (Wong 1993, 1995: Ohmberger 1999; Goh et al. 2010). It is easy to identify

in the field by its leaning-clambering habit, relatively slender culms, and the presence

of conspicuous white hairy bands above and below the culm nodes (Fig. 1).

A recent listing of Thai plant names (Forest Herbarium 2001) had mentioned

the occurrence in Thailand of this bamboo by its scientific name, but without any

specimens cited and without any certain localities indicated. In the Forest Herbarium

(BKF), now under the management of the Department of National Parks, Wildlife and

Plant Conservation, a single specimen consisting only of scanty flowering material and

small leafy branches (C. Nivomdham et al. 2329), collected from Nakhon Si Thammarat,

was located for study. This was not certainly identified, bearing only an annotation that

uses the basionym, “Bambusa cf. montana”. During a bamboo expedition for the Flora

of Thailand Project. specimens of this bamboo (fertile and with all key vegetative parts

including culm leaf sheaths) were collected from the Khao Phanom Bencha National

Park. Krabi province, in Southern (peninsular) Thailand. From this, it has been possible

to identify Maclurochloa montana with certainty. This paper confirms the presence of

M. montana in Thailand, extending the known distribution of this interesting bamboo

further north on the Thai-Malay Peninsula.

298 Gard. Bull. Singapore 62 (2) 2011

a. a. 24/3 .. as

Fig. 1. Recently documented Thai material of Maclurochloa montana. A. Habit and habitat; B.

Details of the culm node and branch complement. Note the characteristic band of white hair on

both sides of the culm node.

Maclurochloa montana in Thailand 299

Additional information based on the Thai material

A description of the species is given in Wong (1995). In addition to that, we can add

the following details:

(1) The culms are 1—3 cm in diameter, with relatively thin walls, 0.2-0.5 cm thick:

with internodes 20—50 cm long.

(2) The culm-sheath has a linear-lanceolate blade: fragile crescentic to lobe-like

auricles 3-5 mm long and 1-2 mm wide, with pale brown bristles 0.5—1 cm long

fringing the margins.

(3) The leaf ligule is a short denticulate to sub-entire rim c. 0.5 mm high, fringed with

a few stiff bristles c. 1-2 mm long, or glabrate.

(4) The pseudospikelet prophylls are 2-keeled, c. 1 mm long by 2 mm wide, with ciliate

keels; the empty glumes are 5—12-nerved; the rachilla internode between perfect florets

is 1—-1.5 mm long, whereas the rachilla internode between perfect floret and terminal

vestigial floret is usually longer, 2—3 mm long; the lemmas are 10—13-nerved; the

paleas are 3—5-nerved between keels and 2—3-nerved between each keel and margin;

the terminal vestigial floret(s) is/are represented only by a lemma, as long as or slightly

exceeding the more distal floret(s), or represented by a lemma and a much reduced

palea; the 3 lodicules are 6—12-nerved, hyaline, ciliate on the margin; the anthers are

3.54.5 mm long, yellow, with acute to pointed tips; the caryopsis is 34 mm long,

bottled-shaped and grooved on one side. (See Fig. 2.)

Fig. 2. Some details of the pseudospikelet and floret of Maclurochloa montana. A. Part of

flowering branch, showing clusters of pseudospikelets. B. A single pseudospikelet, with

prophyll attached basally. C. A dissected pseudospikelet (from left to right: a prophyll; 2 bracts

subtending buds, each followed by its bud: 3 empty glumes; a portion of the rachilla: a perfect

floret and 2 terminal vestigial florets (the uppermost one enfolded by the lower one). D. 2

glumes (left and centre) and a lemma (right) with a reduced palea. E. Lodicule complement. F.

Pistil with 3 stigmas (2 broken). G. A caryopsis. All from C. Rattamanee M234.

300 Gard. Bull. Singapore 62 (2) 2011

Material examined. THAILAND. Southern (peninsular) Thailand: Krabi, Khao

Phanom Bencha National Park, fertile, 16 Jan 2010, C. Rattamanee M234 (BK; BKF:

SING; Herbarium of Faculty of Forestry, Kasetsart University); Nakhon Si Thammarat,

Khao Luang, fertile, 25 Feb 1991, C. Nivomdham et al. 2329 (BKF).

Distribution: Southern (peninsular) Thailand (Krabi and Nakhon Si Thammarat

provinces) to Peninsular Malaysia.

Ecology: Disturbed open sites adjacent to lower montane forest, dominated by grasses

mixed with Me/astoma sp. at 1000-1200 m (C. Rattamanee M234) or montane forest

1500 m above sea level (C. Nivomdham et al. 2329).

Vernacular name: Pai Lueai (That).

Notes: The phylogenetic relationships among various Southeast Asian climbing

bamboo genera, including Maclurochloa, and the core Bambusinae genera Bambusa,

_Dendrocalamus Nees and Gigantochloa Kurz (“BDG complex’), were investigated by

Goh et al. (2010). Their analyses show Maclurochloa to be firmly embedded within

the BDG complex, and the authors discussed the possibility of a reticulate origin for

such taxa although they emphasised the need for more work to clearly demonstrate

this.

ACKNOWLEDGEMENTS. This work was supported by the Commission on Higher Education,

Ministry of Education, Thailand. We thank Dr K.M. Wong, Singapore Botanic Gardens, for

assistance in verifying this bamboo species.

References

Forest Herbarium, Royal Forest Department. (2001) Thai Plant Names: Tem Smitinand

(revised edition). Bangkok: Prachachon Ltd.

Goh, W.L., Chandran, S., Lin, R.-S., Xia, N.-H. & Wong, K.M. (2010) Phylogenetic

relationships among Southeast Asian climbing bamboos (Poaceae: Bambusoideae)

and the Bambusa complex. Biochem. Syst. Ecol. 38: 764-773.

Ohrnberger, D. (1999) The Bamboos of the World: Annotated Nomenclature and

Literature of the Species and the Higher and Lower Taxa. Amsterdam: Elsevier

Science B.V.

Wong, K.M. (1993) Four new genera of bamboos (Gramineae: Bambusoideae) from

Malaysia. Kew Bull. 48(3): 517-532.

Wong, K.M. (1995) The Bamboos of Peninsular Malaysia. Malayan Forest Records

No. 41. Kuala Lumpur: Forest Research Institute Malaysia.

Wong, K.M. (2005) Mullerochloa, a new genus of bamboo (Poaceae: Bambusoideae)

from north-east Australia and notes on the circumscription of Bambusa. Blumea

50: 425-441.

Gardens’ Bulletin Singapore 62 (2): 301-306. 2011 301

The nomenclature of Uvaria velutina Roxb. ex Blume

(Annonaceae)

J.F. Veldkamp

National Center for Biodiversity — Naturalis,

section National Herbarium of The Netherlands, Leiden University

P.O. Box 9514, 2300 RA Leiden, The Netherlands

veldkamp(@nhn.leidenuniv.nl

ABSTRACT. The nomenclature of Uvaria velutina Roxb. ex Blume (Annonaceae) is clarified.

Keywords. Annonaceae, Cananga, Miliusa, nomenclature, SE Asia, Trigynaea, typification,

Uvaria

Introduction

When trying to solve the nomenclature of Uvaria velutina Roxb. ex Blume

(Annonaceae), a seemingly simple problem turned out to be quite complicated and

to involve four species of four different genera: Cananga odorata (Lam.) Hook.f. &

Thoms., Guatteria australis A.St.-Hil., Miliusa velutina (DC.) Hook.f. & Thoms., and

Uvaria hirsuta Jack.

Uvaria velutina Roxb. ex Blume

Blume (1825) described a collection from Kuripan, W. Java, Indonesia, as Uvaria

velutina ? Roxb. Very few names in the Bijdragen have a reference to an earlier

publication or author and Uvaria velutina is one of the exceptions. From a survey of

the first 20 pages it became clear that most of the taxa described there were indeed new

and are to be attributed to Blume, but a few taxa had been published previously by e.g.

Linnaeus (1753), Dunal (1817), and De Candolle (1817). It 1s occasionally cited as a

synonym in later literature, e.g. as “Uvaria velutina Blume non Roxb.” (Hooker f. &

Thomson 1855) or as “Uvaria velutina Roxb. ex Blume” (Meade & Parnell 2002). In

a fairly recent checklist of the Asiatic-Australian species of Annonaceae Kefiler et al.

(1995: 87) cited Uvaria velutina thrice with the authorities as Blume (1825; “Roxb. ex

Blume”), Dunal (1817), and Roxb., the last, however, without any further data.

Where had Blume found this name? The Annonaceae had been more or less

simultaneously studied by Dunal and De Candolle in 1817. The monograph of the first

appeared between August and November and the treatment by the latter in the first half

of November. Their descriptions are often nearly the same and references are made by

one to the other. In both publications the name Uvaria velutina is used, with Uvaria

villosa Roxb. as synonym, a name found in Roxburgh (1814) for a plant accessed in

Calcutta in 1796 from Bengal. Dunal (p. 91) noted for his specimen: “DC. v. s. h.

Lamb.”, that is ‘“‘ De Candolle has seen a dried specimen in the Herbarium of Lambert”

and at the end of the description “DC. Syst. univ. ined.”, apparently a provisional

Zz ‘ard. Bull. Singapore 62 (2) 20

302 Gard. Bull. Singar 2 (2) 2011

title for what was shortly afterwards to be published as “Regni vegetabilis systema

naturale”. There De Candolle referred to “Dunal, Monogr. p. 91”. These citations

support the idea that Dunal has the priority, and as he apparently had not seen any

specimen himself, his information of the species must have come from De Candolle

who had. In 1824 De Candolle himself attributed the combination to Dunal, and cited

his own work as a literature reference. I think the correct citation should be Uvaria

velutina DC. in Dunal.

The provenance was given as India orientalis.

It is known that Lambert had received “several thousands of species” directly or

indirectly from Roxburgh between 1796 and 1810. The specimen that De Candolle had

seen must have been one of these and presumably would now be in G (Miller, 1970).

However, it was not included in the IDC microfiche set of the specimens used for the

Prodromus (1824; G-DC) and Dr. P.J.M. Maas (WAG) told me that he had never seen

a Lambert Annonaceae in G, but had so in BM.

There 1s no Uvaria velutina in Roxburgh’s survey of the plants growing in the

Calcutta Botanic Garden (1814: 43, 94), and the names listed in it are rarely provided

with a diagnosis or a reference to one. Uvaria villosa Roxb. is mentioned but all that

-was said was that the plant had been obtained in 1796 from Bengal. A validating

description was published posthumously in 1832. So this could not have been used by

Blume to identify his specimen and as a source of the epithet.

It seems to me that Uvaria velutina Roxb. ex Blume is an etymological hybrid

between Uvaria velutina (Dunal) and Uvaria villosa Roxb. and was not intended as a

“new” species.

In his introduction Blume (1825) complained about the scarcity of scientific

literature in faraway Buitenzorg (Bogor, Java). In view of the rarity of Dunal’s paper,

it seems more likely that Blume had access to either De Candolle (1817) or (1824).

However, in his lengthy paper of 1830 after his return to Europe he cited all three.

Apparently Blume soon realised this misapplication, for in 1830 (p. 31) he

corrected the authorship to Dunal and cited “U. villosa Roxb. mss”. He stated that

the name had been applied to a plant from Bengal, which clearly is a reference to

Roxburgh (1814). He seems to have been the first after the latter to have given this

provenance; the references he cited all stated “India orientalis”. Curiously, he retained

the name for a fragmentary collection by Reinwardt consisting of a branch and some

leaves, either from Java or the Moluccas (L sh. 898.63--533). By the way, from this it

can be observed that Blume used the Rijnland inch (2.616 cm), although in 1809 that

had officially been decreed to be | cm long.

Miquel (1858) regarded the Reinwardt specimen as belonging to Cananga

odorata (Lam.) Hook.f. & Thoms., yet in 1865 described it as new, Uvaria subcordata.

The first identification was supported by Boerlage in 1899 on the label of the isotype in

BO (two leaves and a satchel with the rather typical shrivelled young branch tips). The

name was cited but the sheet not labelled by Koorders & Valeton (1903) when they

proposed the combination Canangium odoratum var. velutinum [Blume] Koord. &

Valeton. (The square brackets here are used to indicate that there is something “wrong”

with the basionym. This useful method of citation was deleted from the ICBN after

1972). It was confirmed again by Turner who saw the specimen in 2008 (pers. comm.),

and also by KeBler on the holotype in L (det. slip, 2000).

Blume (1830: 22, t. 5) described and depicted his “own” Uvaria velutina from

Kuripan as Uvaria hirsuta.

Nomenclature of Uvaria velutina 303

However, confusion still reigned, for unknown to him there was already a

Uvaria hirsuta by Jack (1820) and another by Vellozo (1829).

Later authors have regarded Jack’s and Blume’s names as taxonomic synonyms,

e.g. Wallich (1832) (followed by Steudel, 1841), Hooker f. & Thomson (1855; 1872),

Miquel (1858), Sinclair (1955), and Backer & Bakhuizen f. (1964). Jack’s name was

neotypified by Meade & Parnell (2002), who did include “Uvaria velutina Roxb. ex

Blume”, but not Blume’s use of Uvaria hirsuta for it. They thought that the Blume

collection would be in BO, but his main set is in L with duplicates in many other

institutes (see Van Steenis-Kruseman, 1950), and some have been distributed to BO.

Guatteria velutina A.DC. (1832) has been regarded as a new combination based

on Uvaria velutina DC. by e.g. Hooker f. & Thomson (1855) and KeBler et al. (1995:

30). However, it is clear from his text that A. De Candolle intended this as a new

species, as he gave no reference to an earlier publication on which this would be based,

but included it in a paragraph entitled “Species Prodromo addendae” (Species that

must be added to the Prodromus). He also gave the provenance as Burma, near the

Atran River, not Bengal. No mention of the taxon was made under Uvaria.

Steudel (1841) proposed Uvaria blumeana as a new name for Uvaria hirsuta

Blume (and of Jack, according to Wallich), and Uvaria velutina Blume (non “Dec.”,

i.e. DC.), as he erroneously thought that Uvaria hirsuta would be a later homonym of

“Arrab.”, 1.e., Vellozo (1829). Uvaria hirsuta Jack and Uvaria hirsuta Vell. are earlier,

valid and legitimate names though, so he created a superfluous one, applicable to the

oldest combination by Jack, although he named it after Blume.

Vellozo’s name is a synonym of Guatteria australis A.St.-Hil. (Maas, 1n litt.),

and not of Trigynaea oblongifolia Schltdl., as was said by Meade & Parnell (2002).

Meade & Parnell (2002) also thought that the type of Uvaria trichomalla Blume

would be a Blume collection in BO, but in fact it is Kuhl & Van Hasselt 1837 in L. This

was seen and identified by Meade himself (det. slip 1/7/2000) as U. hirsuta Jack, but

no L specimen is accounted for in the Meade & Parnell paper.

Many authors have included Uvaria pilosa Roxb. (1832), aname probably based

on a plant cultivated in the Calcutta Botanic Garden as a “native of the Moluccas”,

where Uvaria hirsuta does not occur. This is another example of a mistaken origin

by Roxburgh probably caused by his connections with Christopher Smith. The latter

used to work as the Superintendent of the Botanic Gardens in the Moluccas in Ambon

(1796-1802), and later (1805—1806) with the same rank in Penang, Malay Peninsula.

From Penang he sent plants to Roxburgh, who then thought they had come from the

Moluccas.

Kurz (1877) has reported Uvaria hirsuta from Pegu (Bago), S. Burma, which

was copied by later authors, but not by Kress et al. (2003) who instead mentioned Mon

and Taninthayi (Tenasserim). Ban (2000) reported it for S Vietnam [Quang Binh (Vinh

Linh), Quang Tri (Zakhtajan 212, LE), and Da Nang (Tourane: Poilane 8068, HM, P)].

If this species really occurs in Burma and Vietnam, it would be expected to occur in

Thailand and Cambodia as well. Meade (2000) did not report on what Kurz had seen,

presumably now in CAL. The Vietnam specimens cited by Ban I have not found in his

thesis, either.

However, Meade did not include Uvaria hirsuta and in Meade & Pennington

(2002) stated: “failed to locate a single specimen or reference to a specimen of Uvaria

hirsuta or its synonyms from north of the Thai-Malay border’. This is confirmed by

the experiences of Dr. P. Chalernglin (Bangkok), Mr. J.F. Maxwell (CMU), and Ms. P.

Sidisunthorn (BKF).

304 Gard. Bull. Singapore 62 (2) 2011

From these data it would seem that these records are based on misidentifications.

In the Malaysian Peninsula the species is widespread but does not seem to reach

the Thai / Malay border (Turner 1997), and furthermore occurs in Singapore, Sumatra

(Aceh, West), Bangka, and W Java.

Uvaria hirsuta Jack

Uvaria hirsuta Jack, Malayan Misc. | (1820) 46. — Uvaria blumeana Steud., Nomencl.

2 (1841) 737, nom. superfl. TYPE: Peninsular Malaysia, Penang, lost in the fire of the

“Fame”, 1824, Jack s.n. NEOTYPE: Peninsular Malaysia, Penang, Wallich Cat. 6458-

B (K, holo, IDC microfiche 7394), designated by Meade & Parnell (2002).

Uvaria velutina Roxb. ex Blume, Bijdr. 1 (1825) 13, non Dunal (1817). — Uvaria

hirsuta Blume, FI. Jav. Anon. (1830) 22, t. 5, non Jack (1820), nec Vell. (1829). TYPE:

W. Java, Kuripan, Herb. Blume s.n. (L, sh. 898.63—384).

- Uvaria trichomalla Blume, Fl. Java. Anon. (1830) 42, t. 18. TYPE: W Java, Tjikao,

Kuhl & Van Hasselt 1837 (L, holo).

Guatteria pilosa Roxb. ex G.Don, Gen. Hist. 1 (1831) 98. — Uvaria pilosa (Roxb. ex

G.Don) Roxb., Fl. Ind. 2 (1832) 665. LECTOTYPE: “A native of the Moluccas”, quid

non, Roxburgh s.n. in Wallich Cat. 6458-A (K, holo, IDC microfiche 7394), designated

here.

= Uvaria hirsuta Jack

Uvaria subcordata Miq.

Uvaria ? subcordata Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 9. — Canangium

odoratum var. velutinum {[Blume] Koord. & Valeton, Meded. Lands Plantentuin

61 (1903) 282 [= Biydr. Boomsoort. Java 9 (1903) 282; “velutina’]. TYPE: “Java,

Moluccas”, Reinwardt s.n. (L, holo, sh. 898.63—533; BO, fragm., sh. 1455792).

EPITYPE: Java, Pekalongan, Soebah, Koorders 22570 (L, holo; iso BO), designated

here.

= Cananga odorata (Lam.) Hook.f. & Thoms.

Uvaria velutina DC.

Uvaria velutina DC. in Dunal, Monogr. (1817) 91; DC., Veg. Syst. 1 (1817) 484;

Prodr. 1 (1824) 88. — Guatteria villosa G. Don, Gen. Hist. 1 (1831) 100, nom. superfl.

— Uvaria villosa Roxb., Hort. Beng. (1814) 43, nom. nud.; Roxb., Fl. Ind. 2 (1832)

664, nom. superfl. — Miliusa velutina (DC.) Hook.f. & Thoms., Fl. Ind. 1 (1855) 151.

TYPE: Cultivated in the Botanical Garden of Calcutta, India, Roxburgh s.n. in Herb.

Lambert (? BM, ? G).

Nomenclature of Uvaria velutina 305

Guatteria velutina A.DC., Mém. Anon. (1832) 42. TYPE: Burma, Mon, near the Atran

River, Wallich Cat. 6441-C (K, holo, IDC microfiche 7394).

= Miliusa velutina (DC.) Hook.f. & Thoms.

ACKNOWLEDGEMENTS. Ms. L.-L. Zhou (HK) kindly informed me about some specimens

she had on loan from L. Dr. N.H. Nghia (Forest Science Institute of Vietnam, Hanoi) graciously

translated the description by Ban (2000). Mr. A. Sumadijaya searched for specimens in BO. Dr.

I.M. Turner (K) is thanked for pointing out the problem and checking out material in BO and K.

Dr. P.J.M. Maas (WAG) kindly informed me about the possible location of Lambert collections

of Annonaceae in BM, not G, De Candolle’s role in the Dunal paper, and the present identity

of Uvaria hirsuta Vell. Ms. P. Sidisunthorn (BKF) and Dr. P. Chalernglin (Thailand Institute

of Scientific and Technological Research, Bangkok) advised on the (non-) occurrence of this

species in Thailand. Dr. P.J.A. KeBler (L) assisted in a number of minor questions involving

this paper.

References

Backer, C.A. & Bakhuizen f., R.C. (1964) Flora of Java 1: 103. Groningen: Noordhoff.

Ban, N.T. (2000) Flora of Vietnam. 1. Annonaceae Juss.: 66. Ha Noi: Science &

Technics Publishing House.

Blume, C.L. (1825) Bijdragen tot de flora van Nederlandsch Indié 1: 13 Batavia:

Lands Drukker.

Blume, C.L. (1830) Flora Javae. Anonaceae: 22, 31, t. 5. Brussels: Frank.

De Candolle, A.P. (1817) (1-15 Nov). Regni vegetabilis systema naturale 1: 484.

Strasbourg, London: Treuttel & Wiirtz.

De Candolle, A.P. (1824) Prodromus systematis naturalis regni vegetabilis 1: 88.

Strasbourg, London: Treuttel & Wiirtz.

Dunal, M.-F. (1817) (Aug—Nov). Monographie de la famille des Anonacées: 91-92.

Paris, London, Strasbourg: Treuttel & Wiirtz & Montpellier: Renaud.

Forman, L.L. (1997) Notes concerning the typification of names of William Roxburgh’s

species of Phanerogams. Kew Bull. 52: 513-534 (533).

Hooker, J.D. & Thomson, T. (1855) Flora Indica 1. London: Pamplin.

Hooker, J.D. & Thomson, T. (1872) The Flora of British India 1: 48. Brook nr. Ashford:

Reeve & Co.

Jack, W. (1820) Descriptions of Malayan plants. Malayan Miscellanies 1: 46; reprinted

in Hook., Bot. Misc. 2: 87. 1831.

KeBler, P.J.A., Jessup, L.W. & Kruijer, J.D. (1995) Provisional Checklist of the

Asiatic-Australian Species of Annonaceae: 30, 87. Serdang: The Herbarium, UPM

& UNESCO-Botany Asia 2000.

Koorders, S.H. & Valeton, T. (1903) Bijdrage no. 9 tot de kennis der boomsoorten van

Java. Meded. Lands Plantentuin 61: 282.

Kress, W.J., Defilipps, R., Farr, E. & Daw Yin Yin Kyi (2003) A Checklist of the Trees,

Shrubs, Herbs, and Climbers of Myanmar. Contrib. US Natl. Herb. 45: 144.

Kurz, S. 1877. Forest Flora of Burma 1: 28. Calcutta: Office of the Superintendent of

Government Printing.

306 Gard. Bull. Singapore 62 (2) 2011

Linnaeus, C. (1753) Species Plantarum. Stockholm: Salvius.

Meade , C.V. (2000) A Systematic Revision of the Uvaria L. Group (Annonaceae)

in Continental Asia. PhD Thesis, University of Dublin, Ireland. Ineffective

publication, copy in TCD, L, etc.

Meade, C.V & Parnell, J.A.N. (2002) Neotypification of Uvaria hirsuta Jack

(Annonaceae). Zaxon 51: 767-768.

Miller, H.P. (1970) The herbarium of Aylmer Bourke Lambert. Taxon 19: 538.

Miquel, F.A.W. (1858) Flora van Nederlandsch Indié 1, 2: 24, 40. Leipzig: Fleischer.

Miquel, F.A.W. (1865) Anonaceae archipelagi indici. Annales musei botanici lugduno-

batavi 2: 9. http://www.botanicus.org/item/3 1753002764857.

Roxburgh, W. (1814) Hortus Bengalensis: 43. Serampore: Mission Press.

Roxburgh, W. (1832) (14 Jan) Flora Indica (Carey ed.) 2: 664. Calcutta & Parbury:

Thacker & Co., London: Allen & Co.

Sinclair, J. (1955) A revision of Malayan Annonaceae. Gard. Bull. Singapore 14: 203.

Steudel, E.G. von (1841) Nomenclator Botanicus 2: 737. Stuttgart, Tiibingen: Cotta.

Turner, I.M. (1997) (“1995”). A catalogue of the vascular plants of Malaya. Gard.

Bull. Singapore 47: 121.

- Van Steenis-Kruseman, M.J. (1950) Malaysian plant collectors and collections

(“Cyclopaedia of collectors”). F/. Males. I, 1: 64. Jakarta: Noordhoff-Kol ff.

Vellozo, J.M. de Concei¢ao (1829) Florae Fluminensis: 238, t. 124. Rio de Janeiro:

Typographia Nacionalis.

Wallich, N. (1832) 4 Numerical List of Dried Specimens of Plants in the East India

Company s Museum: Collected under the Superintendence of Dr. Wallich of the

Company 8s: # 6441, 6458. London: Wallich.

Gardens ’ Bulletin Singapore 62 (2): 307-311. 2011 307

Steenisia (Rubiaceae) newly recorded for mainland Asia

with a new variety of S. pleurocarpa (Airy Shaw) Bakh.f.

K.M. Wong

Singapore Botanic Gardens, | Cluny Road, Singapore 259569

wkm2000@gmail.com

ABSTRACT. Mussaendopsis malayana T.Yamaz. (Rubiaceae), described based on material

from the Malay Peninsula, is a distinct variety of Steenisia pleurocarpa (Airy Shaw) Bakh-f.

It represents a new record for mainland Asia of Steenisia Bakh.f., a genus otherwise endemic

to Borneo and the Natuna islands. The new combination Steenisia pleurocarpa var. malayana

(T. Yamaz.) K.M.Wong is made.

Keywords. Asia, biogeography, Borneo, Malay Peninsula, Mussaendopsis, Rubiaceae. South

China Sea, Steenisia, vicariance

Introduction

Mussaendopsis malayana T.Yamaz. (Rubiaceae) was described based on a collection

from around 700 m elevation on the west side of Mount Tahan, between Sungai

Luis and the summit, in Peninsular Malaysia (Yamazaki 2001). A good photograph

of the holotype specimen (Ohba & Miyamoto 90504) and diagnostic drawings of a

vegetative node with a stipule, the axillary inflorescences, flowers and fruits were

provided with the description. Images of the holotype, an isotype sheet, and two sheets

comprising a paratype collection (Ohba & Miyamoto 90292) were made available

from the University of Tokyo herbarium (T1).

These collections from Mount Tahan are identifiable with Steenisia pleurocarpa

(Airy Shaw) Bakh.f. from northwest Borneo, particularly in the very similar habit (low

treelet); leaf form, venation and pubescence; interpetiolar stipules fringed with linear

teeth; paired axillary compound-thyrsoid inflorescences; development of an enlarged

calyx member (semaphyll) in some post-anthesis flowers and fruits; and septicidally

dehiscent fruits with conspicuous ribbing on the outside (Bremer 1984). Yamazaki

(2001) diagnosed the stipules in the Tahan material as intrapetiolar but this is clearly

in error as revealed in his Fig. 2a and the material he cited.

Mussaendopsis and Steenisia compared

Although Mussaendopsis Baill. also has axillary inflorescences in which some flowers

also develop an enlarged calyx member, that genus has many different attributes

compared to Steenisia Bakh.f. Table 1 compares the characters of Mussaendopsis and

Steenisia.

308 Gard. Bull. Singapore 62 (2) 2011

Mussaendopsis is a distinct genus of much taller trees, easily diagnosed by

intrapetiolar stipules, glabrous leaves, right-contorted corolla lobes, free anthers in

the open flower, and seeds that are winged all around (Bremekamp 1939, Puff &

Igersheim 1994). The molecular evidence suggests a placement for Mussaendopsis

within the tribe Condamineeae (Alejandro et al. 2005, Bremer 2009). Steenisia, on the

other hand, is a genus of low treelets and subshrubs (Bremer 1984) with interpetiolar

stipules, pubescent leaves, left-contorted (rarely valvate) corolla lobes, connate

anthers in the open flower, and unwinged seeds that are or only slightly winged at

two ends. There is as yet no indication of the systematic position of Steenisia from

molecular studies (Bremer 2009), although Bremer (1984) had compared it with the

Rondeletieae. It is interesting to note that two other genera with connate anthers in the

open flower, Argostemma Wall. and Neurocalyx Hook.f., once accepted in the same

tribe as Steenisia (Verdcourt 1958), are now placed by molecular inference in the

Argostemmateae and Ophiorrhizeae, respectively (Bremer & Manen 2000, Robbrecht

& Manen 2006, Rydin et al. 2009).

When describing his “M. malayana” Yamazaki (2001), however, had stated

“this is the fourth species in the genus”. He was apparently unaware that one of

“the “Mussaendopsis” species was transferred to Greeniopsis as early as 1909:

Mussaendopsis multiflora Elmer, Leafl. Philipp. Bot. 1: 14 (1906) is now Greeniopsis

multiflora (Elmer) Merr., Philipp. J. Sci., C 4: 325 (1909) (C.Puff, pers. comm.).

The studies of Puff & Igersheim (1994) reveal that Mussaendopsis beccariana

stipules at inception are already fused up to near their apices, forming an appressed,

resin-filled compartment around the young shoot apex, and the position of the apical

cleft in the largely fused structure suggests an interpetiolar position at inception. Later

ontogenetic development of the full stipular structure, however, is clearly intrapetiolar,

with a thickened suture along the interpetiolar median, at which the stipular sheath

splits into two clearly intrapetiolar members. In Steenisia, the apical buds are not

protected in this way, and the stipules are only slightly connate basally and distinctly

interpetiolar throughout.

The development of semaphylls is also different. In Mussaendopsis, these have

already formed as enlarged petaloid structures even at the floral bud stage, although in

those species of Steenisia where these are found, they form only in the post-anthesis

stage. In both cases they persist into the fruit stage.

Professor Christian Puff (pers. comm.) points out that Steenisia (with its anther

cones) is a typical buzz-pollinated taxon (where pollen is vibrated out of the anther

cone by the close buzzing of bees and then collected) (Buchman, 1983), and thus no

nectar is produced as reward. On the other hand, Mussaendopsis 1s fly-pollinated: the

rotate corolla allows good landing space and a huge fleshy, disc-like nectar-producing

structure surrounds the base of the style (the flowers resembling those of numerous

Celastraceae taxa).

A new variety of Steenisia pleurocarpa

Bremer (1984) distinguished the various Steenisia species by characters such as crowded

or well-spaced leaf arrangement along the stem, shape of the leaf base, calyx shape,

presence / absence of semaphylls, semaphyll form and venation, style pubescence,

ribbing of the fruit pericarp, and seed shape. The Malayan taxon is identifiable to

Steenisia new for mainland Asia

Table 1. Differentiating characteristics of Mussaendopsis and Steenisia.

Leaf hairiness

Stipule insertion

Stipule margin

Mussaendopsis

(2 spp. known)

glabrous

intrapetiolar, connate for nearly

whole length at first

following separation, margins

entire except for tiny apical

Steenisia

(5 spp. known)

pubescent

interpetiolar, only

basally connate

laciniate to fringed

with linear teeth

cleft

Corolla lobe aestivation night-contorted left-contorted, rarely

valvate

Anthers in the open widely spaced out connate into a central

flower cone-like structure

Seeds winged all around not winged, or very

slight wings at

opposite ends only

Enlargement of one

calyx lobe in a flower /

fruit as a semaphyll

semaphyll present from flower

bud stage

semaphyll developing

at post-anthesis

Steenisia pleurocarpa, being identical in all these characteristics, overall habit (as low

treelets), leaf form, and fruit form and size. Mature corollas were not available for the

Malayan collections and so could not be directly compared, but flower bud material

did not suggest any potential differences.

The only differences between the Malayan taxon and the Bornean material

of Steenisia pleurocarpa are in peduncle length and degree of branching of the

infructescence. Fourteen infructescences were measured for Bornean material, with

peduncle length ranging 5.4-8.7 cm long, and the two infructescences in the Malayan

material from Mount Tahan had peduncles that were 1.5 and 1.6 cm long. Also, the

Bornean material had 3-4 (rarely 2) pairs of branches on the infructescence rachis,

with the lowest branches ramifying to 3—5 orders (rarely 2 orders). In the Malayan

material, the infructescences had only 1-2 pairs of branches on the infructescence

rachis, with just 1—2 orders of ramification. Therefore, the Malayan material appears

to have distinctly shorter, less branched infructescences compared to the Bornean

material.

Here we propose to consider the Bornean and Malayan material as distinct

varieties. The Malayan material is named as follows.

310 Gard. Bull. Singapore 62 (2) 2011

Steenisia pleurocarpa (Airy Shaw) Bakh.f. var. malayana (T.Yamaz.) K.M.Wong,

comb. et stat. nov.

Basionym: Mussaendopsis malayana T. Yamaz., J. Jap. Bot. 76 (2001) 28-30. TYPE:

Ohba & Miyamoto 90504, Peninsular Malaysia, Pahang, Taman Negara, west side

of Gunung Tahan, Sungai Luis — Summit, 700 m alt., 102°05’E 4°41’N to 102°15’E

4°39’N, 18 Mar 1990 (holotype TI, isotype TI).

Biogeographic implications

Steenisia was previously known only on Borneo (five species) and the Natuna islands

(S. borneensis (Valeton) Bakh.f., one of the five mentioned). With the diagnosis of

Steenisia pleurocarpa var. malayana, the genus 1s now newly recorded for the Malay

Peninsula and mainland Asia. The genus thus has a distribution spanning both sides

of the present South China Sea. The differences between both varieties of Steenisia

pleurocarpa could be explained by vicariance of a formerly widespread ancestral

taxon in this area, separating into two isolated populations that began to diverge

~ evolutionarily.

This is plausible because there is palaeo-ecological, geomorphological and

fossil evidence indicating periods of dryland contiguity between the Malay Peninsula

and Borneo during the Quaternary from about 1.8 Ma to the Last Glacial Maximum

(Flenley 1979, 1984; Morley & Flenley 1987; Stuijts et al. 1988; Morley 2000), when

intervening sea levels could have fallen as much as 40—120 m below current level,

allowing vegetation development and significant species range extensions (Heaney

1991, Voris 2000). These periods alternated with others during which sea-level rises

would have segregated populations of organisms.

The occurrence of Steenisia borneensis in both Borneo and the Natuna islands in

the South China Sea probably also bespeaks such former land connections that are now

interrupted by present sea levels. Similarly, the gecko Ptychozoon lionotum, monitor

lizard Varanus nebulosus and snake Trimeresurus puniceus are found in the Malay

Peninsula with their eastern limit at the Natunas, and some Bornean herpetofauna also

reach westward only as far as the Anambas and Natuna islands (Leong et al. 2003).

ACKNOWLEDGEMENTS. Dr K.T. Yong (KLU) kindly arranged for a copy of Yamazaki’s

paper to be made available through the British Library. I thank Dr Benito Tan and Serena Lee

of the Singapore Herbarium (SING), the Tokyo Herbarium (TI) and Professor Jin Murata of

the Graduate School of Science, University of Tokyo, for arranging for images of the type

material for this study. Professor Christian Puff (University of Vienna) gave useful comments

that improved the paper.

References

Alejandro, G.D., Razafimandimbison, S.G. & Liede-Schumann, S. (2005) Polyphyly

of Mussaenda inferred from ITS and trnT-F data and its implication for generic

limits in Mussaendeae (Rubiaceae). Amer. J. Bot. 92: 544-557.

Steenisia new for mainland Asia 311

Bremekamp, C.E.B. (1939) The genus Mussaendopsis Baill. (Rub.). Recueil Trav. Bot.

Neerl. 36: 367-371.

Bremer, B. (2009) A review of molecular phylogenetic studies of Rubiaceae. Ann.

Missouri Bot. Gard. 96: 4-26.

Bremer, B. & Manen, J.F. (2000) Phylogeny and classification of the subfamily

Rubioideae (Rubiaceae). Plant Syst. Evol. 225: 43-72.

Buchmann, S.L. (1983) Buzz pollination in angiosperms. In: Jones, C.E. & Little, R.J.

(eds) Handbook of Experimental Pollination Biology, pp. 73-113. New York: Van

Nostrand Reinhold.

Flenley, J.R. (1979) The Equatorial Rain Forest: A Geological History. London:

Butterworths.

Flenley, J.R. (1984) Late Quaternary changes of vegetation and climate in the Malesian

mountains. Erdwiss. Forsch. 18: 261-267.

Heaney, L.R. (1991) A synopsis of climatic and vegetational change in Southeast Asia.

Climate Change 19: 53-61.

Leong, T.M., Grismer, L.L. & Mumpuni (2003) Preliminary checklists of the

herpetofauna of the Anambas and Natuna islands (South China Sea). Hamadryad

27: 165-174.

Morley, R.J. (2000) Origin and Evolution of Tropical Rain Forests. Chichester: Wiley.

Morley, R. & Flenley, J.R. (1987) Late Cainozoic vegetational and environmental

changes in the Malay Archipelago. In: Whitmore, T.C. (ed) Biogeographical

Evolution of the Malay Archipelago, Oxford Monographs in Biogeography no. 4,

pp. 50-59. Oxford: Oxford Scientific Publications.

Puff, C. & Igersheim, A. (1994) The character states of Mussaendopsis Baill.

(Rubiaceae — Coptosapelteae). Flora 189: 161-178.

Robbrecht, E. & Manen, J.F. (2006) The major evolutionary lineages of the coffee

family (Rubiaceae, angiosperms). Syst. Geogr. Pl. 76: 85-146.

Rydin, C., Kainulainen, K., Razafimandimbison, S. G., Smedmark, J. E. E. & Bremer,

B. (2009) Deep divergences in the coffee family and the systematic position of

Acranthera. Pl. Syst.Evol. 278: 101-123.

Stuijts, I., Newsome, J.C. & Flenley, J.R. (1988) Evidence of Late Quaternary

vegetation change in the Sumatran and Javan highlands. Rev. Palaeobot. Palynol.

55: 207-216.

Verdcourt, B. (1958) Remarks on the classification of the Rubiaceae. Bull. Jard. Bot.

Etat Bruxelles 28: 209-290.

Voris, H.K. (2000) Maps of Pleistocene sea levels in Southeast Asia: shorelines, river

systems and time durations. J. Biogeogr. 27: 1153-1167.

Yamazaki, T. (2001) A new species of Mussaendopsis (Rubiaceae) from the Malay

Peninsula. J. Jap. Bot. 76: 28-30.

Gardens’ Bulletin Singapore 62 (2): 313-325. 2011 313

Studies on Homalomeneae (Araceae) of Borneo III:

The helophytic Homalomena of Sunda

Sin Yeng Wong’, P.C. Boyce? and B.A. Fasihuddin®

' Department of Plant Science & Environmental Ecology,

Faculty of Resource Science & Technology, Universiti Malaysia Sarawak,

94300 Kota Samarahan, Sarawak, Malaysia

sywong@frst.unimas.my

> Pusat Pengajian Sains Kajihayat (School of Biological Sciences),

Universiti Sains Malaysia, 11800 USM, Pulau Pinang, Malaysia

phymatarum@gmail.com

Faculty of Resource Science & Technology, Universiti Malaysia Sarawak,

94300 Kota Samarahan, Sarawak, Malaysia

ABSTRACT. An account of the helophytic Homalomena in Sunda is presented. Two species

are recognized: H. expedita A.Hay & Hersc. and H. rostrata Griff., neither novel. Homalomena

expedita is so far endemic to Sarawak, where it is known from three widely separated sites.

Homalomena rostrata is widespread from Sumatera through Peninsular Malaysia to Borneo, and

onwards to Maluku (Pulau Ceram). Throughout its range H. rostrata is morphologically plastic,

such that it has accrued a considerable synonymy, proposed here: Homalomena beccariana

Engl., H. ensiformis Alderw., H. miqueliana Schott, H. miqueliana var. truella Alderw., H.

paludosa Hook.f., H. propinqua Schott, H. raapii Engl., H. sagittifolia Jungh. ex Schott,

H. sagittifolia var. angustifolia Furtado, H. sagittifolia var. pontederiifolia Ridl. (including

homotypic H. ridleyana Engl.), H. sagittifolia var. sumatrana Alderw., H. tevsmannii Engl..

and H. triangularis Alderw. An overview of the occurrence of helophytism in Araceae, together

with speculations on the evolution of helophytism in Homalomena, a key to the species, and

illustrations of both species are presented.

Keywords. Araceae, helophyte, helophytism, Homalomena

Introduction

Homalomena in Sarawak is presently the subject of study by a taxonomic and

systematic consortium coordinated from the Universiti Malaysia Sarawak (UNIMAS)

(see Boyce & Wong 2008; Boyce & Wong 2009; Boyce, Wong & Fasihuddin 2010;

Ng et al., in prep.). One of the first priorities has been the application of previously

published names, a task especially problematic in Homalomena due to the poor state of

preservation of most of the historical types. Nonetheless, progress has been made, and

the taxonomic status of most published Bornean Homalomena names is now resolved.

Disturbingly, this process has resulted in just 13 ‘good’ names for Borneo, plus about

10 as yet unresolved, and these virtually all from Kalimantan. Given the very high

levels of local endemism that is a feature of Homalomena, and that Sarawak alone

likely has in excess of 300 species, it can be confidently stated that nearly everything

encountered in the field in Sarawak would be undescribed.

This paper is one in a continuing series that aims to bring a better understanding

to this, the most speciose and complex genus of the Araceae of SE Asia. Of the species

314 Gard. Bull. Singapore 62 (2) 2011

dealt with here, H. rostrata Griff. is one of the very few widespread and also highly

polymorphic species in a genus that is otherwise notable for an exceptionally high

degree of localized endemism and low levels of morphological variation. The large

amount of material of H. rostrata in regional herbaria has been almost universally

misidentified, and in addition the two names most commonly applied to almost all

cordate-leaved Homalomena seen in herbaria, H. sagittifolia Jungh. ex Schott and H.

propinqua Schott, are both junior synonyms of H. rostrata. This paper aims to settle

the taxonomic chaos that exists around this name.

Homalomena expedita A.Hay & Hersc. is included in this paper as the only

other colonially helophytic species of Homalomena and as such is important to include

as it is potentially a source of taxonomic confusion. Furthermore, H. rostrata and H.

expedita belong to phylogenetically widely separated parts of the genus and thus are

certainly an example of independent evolution into the (for Araceae) rare ecology of

colonial helophytism.

The overwhelming majority of Homalomena are terrestrial or lithophytic

clumping mesophytic herbs occurring in shady perhumid or everwet forest. However,

two species, H. expedita and H. rostrata are colonial, stoloniferous helophytes.

’ Whereas the taxonomy of the recently described H. expedita poses no problems, and

the species is remarkably stable in terms of gross morphology, perhaps as a result of its

clonal colonial habit (see below), H. rostrata is exceedingly morphologically variable,

as well as widespread, with the result that it has been redescribed no fewer than 15

times.

Taxonomic Part

Key to helophytic Homalomena in Sunda

Leaves very broadly ovato-sagittate; spathe not constricted; spadix about 4.5 cm

long, with a conspicuous warty interstice between the male and female flower zones,

interpistillar stamunodes aDSent 2itcceee nsec eee eee 1. H. expedita

Leaves sub-linear to broadly ovato-oblong; spathe conspicuously constricted; spadix

up to 14 cm long, male and female flower zones contiguous, interpistillar staminodes

PYCSEMNE «...cs00ccssccscessesesevasctacheaaescecosecnpeec Oee eee ae a 2. H. rostrata

1. Homalomena expedita A.Hay & Hersc., Gard. Bull. Singapore 54: 174 (2002).

TYPE: Cult. RBG Sydney Acc. No. 940562 ex Malaysia, Sarawak, Lundu, near bridge

[over the Batang Kayang] on Kuching Road (orig. coll. Hay, Yahud, Saupel & Chan

9409), C. Herscovitch s.n. (holo NSW, iso SAR!). (Fig. 1 & 2)

Colony-forming stoloniferous strongly aromatic (terpenoids — anethol?) helophytic

herbs to c. 75 cm tall. Stem an erect to creeping rhizome to c. 30 cm long, c. 4 cm thick,

spongy, emitting cataphylliferous stolons to c. 40 cm long, | cm thick, these eventually

upturned, becoming rhizomatous, leafy, and thence emitting further stolons from the

base of the rhizomatous portion. Leaves clustered, up to c. 10 together; petiole to c.

45 cm long, spongy within, pale mid-green with broken darker green longitudinal

316 Gard. Bull. Singapore 62 (2) 2011

striations, sheathing in the lower '/,; petiolar sheath margins mainly in-rolled except

at the base, sheath thus closed; Jamina mid-green on both sides, initially glossy, later

becoming matt, very broadly ovato-sagittate, leathery, c. 15-20 cm long and wide, the

apex broadly obtuse, very abruptly and shortly acuminate for c. 1 cm, finally stiffly

apiculate for c. 3 mm, the base shallowly cordate to almost truncate, usually distinctly

asymmetric, with widely spreading rounded to subtriangular posterior lobes 7-10 cm

long; midrib adaxially flat, abaxially slightly prominent, with c. 5 adaxially impressed

abaxially slightly prominent primary lateral veins on each side (plus a cluster of 2 or

3 on each side running to the posterior lobes), alternating with fainter interprimaries

and diverging at c. 60°. Inflorescence solitary; peduncle 7-20 cm long, rather thick

and spongy, c. 8 mm diam., erect. Spathe green, c. 5 cm long, 1.5 cm across and in

bud slightly inflated at level of female zone, thence subcylindric-tapering, but not

constricted, later very narrowly ovoid, apiculate for c. 4 mm. Spadix 4.5 cm long,

stipitate; stipe c. 6 mm; female flower zone more-or-less cylindric, 0.8—1.2 cm long

(irregular length around circumference of spadix), 1.2 cm wide; pistils subglobose, 1—2

mm diam.; stigmas subsessile, discoid or very weakly lobed, slightly narrower than the

ovary, papillate; infrapistillar staminodes absent, sterile interstice conspicuous, 1.5 cm

~ long, naked, pale green and c. 6 mm diam. in the lower | cm, with more or less regular

spirals of low domed warts c. 0.5 mm diam., the upper part (comprising the base of

the male zone) 9 mm diam., ivory and clothed in irregular sterile stamens: fertile

male zone elongate-bullet-shaped, 1.5 cm long, tapering to a blunt acute tip, ivory;

male flowers irregular (1—)2—4-staminate; stamens truncate, irregular in size, 1—1.5

mm across, irregularly polygonal, slightly sinuous on the abaxial side, the thecae

overtopped by a synconnective. Fruit unknown.

Distribution: Malesia: endemic to Sarawak; there known from the extensive type

locality, a tiny relictual population along Jalan Steven Yong (Kuching/Bau border),

and also sighted near Sibu along the main road to Miri.

Habitat: Forming large, probably clonal colonies in open swamps and ditches at low

elevation, sometimes in tidal mud with Cryptocoryne ciliata (Roxb.) Fisch. ex Wydler,

and mangroves.

Other material seen: SARAWAK. Kuching Division: Lundu, Kampung Stenggang

Melaya Baru, just after the bridge across the Batang Kayang on the Lundu side of the

river, 01°39°03.8"°N 109°51°40.5”E, 3 Mar 2004, P.-C. Boyce & Jeland ak Kisai AR-

211(SAR); 10 Oct 2004, P.C.Boyce & Jipom ak Tisai AR-724 (SAR); 11 May 2008

P.C. Boyce AR-2357 (SAR).

Etymology: The epithet means ‘set free’, or ‘foot-loose’, alluding to the departure from

confinement to shaded conditions otherwise typical for the genus, and the invasive,

stoloniferous habit.

Notes: Homalomena expedita is remarkable in this genus of predominantly mesophytic

shade-loving terrestrial plants for its occupation of open swampy sites, even tidal

mudflats, in full sun. The spadix of Homalomena expedita is unique in the genus,

having a very conspicuous naked warty interstice above the female zone.

Helophytic Homalomena of Sunda 317

Fig. 2. Homalomena expedita A.Hay & Hersc. A. Active shoot with two post anthesis

inflorescences (oldest to the right) and prophyll subtending reiterative axis of active shoot. B.

Inflorescence at female anthesis. Note the numerous Colocasiomyia (Diptera, Drosophilidae).

C. Inflorescence at onset of male anthesis with a solitary beetle (Chrysomelidae: Dercetina?).

The damage to the male portion of the spadix is typical for chrysomelid visited inflorescences.

D. Spadix at female anthesis with spathe artificially removed. Note the distinctive warty texture

of much of the sterile interstice. E. Detail of the sterile interstice. The morphology of the warts

and their transitional morphology through staminodes to functional male flowers suggest that

they are aborted stamens. Photo credits: P.C. Boyce.

318 Gard. Bull. Singapore 62 (2) 2011

Homalomena expedita was described less than a decade ago, based on collections

made in the 1994, although specimens in FI-B seen by the first author reveal that

Odoardo Beccari twice collected this species, in a locality where it is probably now

extinct, in the 1860s, but failed to recognize its novelty, as too did Engler when working

up Beccari’s collections in the 1870s (Boyce & Wong 2009).

Homalomena expedita appears to spread rapidly through the production of

stolons. At Lundu it forms dense swards in shallow muddy ponds and from a distance

rather resembles water hyacinth (Eichhornia: Pontederiaceae). It appears less vigorous,

though nonetheless invasive and persistent in tidal brackish conditions at Sibu (Hay &

Boyce, pers. obs.).

In relation to the rest of its genus, Homalomena expedita is rather analogous to

the open swamp-dwelling, colony-forming Aglaodorum, contrasted with Ag/laonema

(both Aglaonemateae). Both Homalomena expedita and Aglaodorum griffithii Schott,

besides their similar habitat preferences, and spongy rhizomes, petioles and peduncles,

predominantly produce solitary inflorescences where their immediate forest-dwelling,

non-proliferating relatives generally produce complex synflorescences (except in some

- of the most diminutive species). In addition to the reduced number of inflorescences, it

seems, on the basis of limited observations of H. expedita, that there is other evidence

pointing to depressed or suppressed sexual fertility; the papillae on the interstice

appear to be abortive pistils; the staminal thecae contain some pollen, but are partially

empty in most anthers; senescent male flowers appear not to have shed pollen, the

ovaries however, are filled with ovules which appear normal in form.

Inflorescences of H. expedita attract numerous flies of the genus Co/ocasiomyia

(Diptera, Drosophilidae) and also beetles of Chrysomelidae (probably a Dercetina sp).

The latter also damage portions of the male flower zone (see Fig. 2 C). It is not yet

know which, if either, 1s the effective pollinator.

Homalomena expedita is quite often seen in cultivation in Sarawak; it seems

likely that other populations than those listed above exist and are the source of these

cultivated plants. Intriguingly, it is also encountered for sale in markets in Bangkok,

with the sellers consistently claiming that the plants originate from southern Thailand.

Given the widespread range of other Asian helophytic aroids (e.g., Homalomena

rostrata, Cryptocoryne ciliata, Aglaodorum griffithii) 1t is by no means impossible that

H. expedita is much more widespread but simply overlooked; its close resemblance, at

least at a distance, to weedy Pontederiaceae might in part explain this.

2. Homalomena rostrata Griff., Not. Pl. Asiat. 3: 154 (1851) [‘roshalum’ sphalm.].

TYPE: Malaysia, Malacca: W. Griffith 5989 (K, holo!). (Fig. 3)

Cyrtocladon sanguinolentum Griff., Not. Pl. Asiat. 3: 147 (1851). — Chamaecladon

sanguinolentum (Griff.) Schott, Prodr. Syst. Aroid.: 316 (1860). TYPE: Malaysia,

Malacca, W. Griffith 5990 (holo K!).

Homalomena sagittifolia Jungh. ex Schott, Prodr. Syst. Aroid.: 311 (1860), syn. nov.

TYPE: Malaysia, Sabah, Labuan, Motley (holo K!); epitype designated here: Indonesia,

Java, FEW. Junghuhn $s.n. Wt; Schott Ic., no. 2111 (W!). — Fiche no. 44: a4 in the

microfiche edition. See Notes on typifications, below.

Helophytic Homalomena of Sunda 319

Homalomena miqueliana Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 126 (1863).

TYPE: Indonesia, Borneo, Kalimantan. P W. Korthals s.n. (holo L!).

Homalomena propinqua Schott, Ann. Mus. Bot. Lugduno-Batavi 1: 280 (1864). syn.

nov. TYPE: Indonesia, Kalimantan, Kalimantan Selatan, Mt. Gintang, P W. Korthals

s.n. (holo L!).

Homalomena beccariana Engl., Bull. Soc. Tose. Ortic. 4: 296 (1879). syn. nov. TYPE:

Malaysia, Sarawak, Kuching, Jul 1865, O. Beccari PB. 260 (holo FI-B!).

Homalomena paludosa Hook.f., Fl. Brit. India 6: 531 (1893). LECTOTYPE selected

here: Malaysia, Perak, Larut, H.H. Kunstler (Dr King’s Collector) 3082 (lecto K!

isolecto CAL, SING!). See Notes on typifications, below.

Homalomena sagittifolia var. pontederiifolia Ridl., J Straits Branch Roy. Asiat. Soc.

44: 172 (1905) — Homalomena ridlevana Engl., Bot. Jahrb. Syst. 37: 123 (1905).

TYPE: Malaysia, Borneo, Sarawak, Kuching, E./. Bartlett & G.D. Haviland 3134

(holo SING!).

Homalomena teysmannii Engl., Pflanzenr., [V, 23Da: 68 (1912). TYPE: Indonesia,

Sumatera, Bangka Island, JE. Teijsmann 3227 (holo B! iso BO!).

Homalomena raapii Engl. Pflanzenr., IV, 23Da: 73(1912), syn. nov. LECTOTYPE

selected here: Indonesia, Sumatera, H. Raap 235 (lecto BO!). See Notes on typifications,

below.

Homalomena triangularis Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 181 (1922), syn.

nov. TYPE: Indonesia, Sumatera, Deli, J.A. Lérzing 4071, 27 Jul 1915 (holo BO!).

Homalomena sagittifolia var. sumatrana Alderw., Bull. Jard. Bot. Buitenzorg, III, 4:

192 (1922), syn. nov. LECTOTYPE selected here: Indonesia, Sumatera, Ophir, Taloe,

10 Jul 1917 H.A4.B. Biinnemeijer 129, (lecto BO!).

Homalomena ensiformis Alderw., Bull. Jard. Bot. Buitenzorg, II], 4: 335 (1922).

TYPE: Indonesia, Kalimantan, JG. Hallier 1465 (holo BO! iso L!).

Homalomena miqueliana var. truella Alderw., Bull. Jard. Bot. Buitenzorg, III, 4: 336

(1922), syn. nov. TYPE: Indonesia, Sumatera, Riau Archipelago, Ampoelai, Pulau

Bintang, 14 Jun 1919, H_A.B. Biinnemeijer 6200 (holo BO: iso L!).

Homalomena sagittifolia var. angustifolia Furtado, Gard. Bull. Straits Settlem. 10:

228 (1939), syn. nov. LECTOTYPE selected here: Malaysia, Johore, Mt Austin, H.N.

Ridley 12018 (lecto SING! isolecto K!). See Notes on typifications, below.

Clump or colony-forming stoloniferous very strongly aromatic (terpenoids — ocimene

or carene?) usually helophytic herbs to c. 1 m tall. Stem an erect to creeping rhizome

to c. 20 cm long, c. 2 cm thick, somewhat spongy, emitting cataphylliferous stolons

to c. 25 cm long, | cm thick, these eventually upturned, becoming rhizomatous, leafy,

320 Gard. Bull. Singapore 62 (2) 2011

and thence emitting further stolons from the base of the rhizomatous portion. Leaves

clustered, up to c. 20 together; petiole 10—S0 cm long, distally weakly to rather strongly

D-shaped in cross-section , rather spongy within, deep-green to dark red or brown,

especially for the basal half, with conspicuous broken dark green to brown longitudinal

striations, sheathing in the lower '/, to half petiolar sheath broadly winged, the wings

spreading and somewhat fleshy, sheath open, the petiole essentially canaliculate in

cross-section; /amina deep green to deep brown, glossy, very variable in overall shape,

ranging from sub-linear to broadly ovato-oblong, leathery, c. 10-50 cm long, 3—25

cm wide, the apex ranging from acuminate to broadly obtuse, abruptly and shortly

acuminate for c. 1 cm, finally stiffly apiculate for c. 10 mm, base decurrent-cuneate

to truncate, shallowly cordate to sagittate or hastate, usually distinctly asymmetric,

posterior lobes where present straight to rather widely divergent, up to 12 cm long,

somewhat acutely rounded; midrib adaxially flat, abaxially prominent, with c. 5—7

adaxially impressed abaxially rather prominent primary lateral veins on each side,

posterior lobes where present with 2 or 3 clustered veins running into them, primary

lateral veins alternating with somewhat fainter interprimaries and diverging all at c.

- 60° from the mid-rib. Inflorescences 2—3 together, the synflorescence subtended by a

conspicuous prophyll; peduncle 8—15 cm long, 5—10 mm diam., robust and somewhat

spongy, medium green to reddish or brown, especially near the base, exceptionally

glossy bright red. Spathe externally green, flushed red, or more rarely deep red, rather

glossy, internally somewhat pale, rather variable in size, 5—15cm long, conspicuously

constricted at a point corresponding to the top of the female flower zone: opening by

inflation of the lower spathe and loosening of the spathe limb (female anthesis), and

then spreading of the spathe limb (male anthesis); post anthesis spathe closing and

tightly clasping the spadix: lower spathe oblongo-ovate to ovato-globose, c. '/, spathe

length, up to 5 cm long, 2 cm wide, externally with conspicuous g glands (extrafloral

nectaries?); spathe limb narrowly to somewhat broadly triangular, c. */, spathe length,

up to 7 cm long, 3 cm wide, rostrate-apiculate for c. 10 mm. Spadix subequalling the

spathe, up to 14 cm long; shortly stipitate; stipe c. 6 mm; female flower zone weakly to

rather strongly fusiform, up to 5 x 1.5 cm; pistils subglobose, 1—2 mm diam., green to

red depending on the overall colour of the spathe, red pistils associated mainly with red

or red-flushed spathes; stigmas subsessile, discoid to rather well-lobed, slightly wider

than the ovary, occasionally remarkably robust, papillate at anthesis; interpistillar

staminodes weakly to very strongly clavate, subequalling the associated pistil, waxy

white; male flower zone tapering-cylindrical, up to 10 cm long, 1 cm wide, tapering to

an acute tip, ivory to very pale yellow; male flowers irregular 3—4-staminate; stamens

truncate, rather regular in size, 1-2 mm across, polygonal, slightly sinuous on the

abaxial side, the thecae overtopped by a pronounced synconnective. Infructescences

declinate by flexing of the peduncle; spathe persistent, turning deep red; at fruit maturity

spathe shed by the abscising of the lower spathe at the insertion on the peduncle. Fruit

oblong-globose, dull red, smelling strongly of butyric acid. Seeds elongate-ellipsoid,

c. 1 x 0.3 mm, very finely longitudinally striate, pale brown.

Distribution: Malesia: S Peninsular Thailand (Narathiwat), Sumatera, Peninsular

Malaysia, Borneo, Maluku (Pulau Ceram).

Helophytic Homalomena of Sunda

Fig. 3. Homalomena rostrata Griff. A & B. Plants in habitat showing variation in leaf lamina

shape. C. Sub-mature infructescence. D. Mature infructescence with spathe artificially

removed. Note the glands on the lower spathe exterior. Images: A: AR-/416; B: AR-1372; C:

AR-2600; Photo credits: P.C. Boyce. D: S. Lee 3/9. Courtesy of Singapore Botanic Gardens,

used with permission.

B22, Gard. Bull. Singapore 62 (2) 2011

Habitat: Swampy areas in lowland forest, forest margins, in freshwater or peatswamp,

or wet facies of kerangas, or kerapah, often, but not exclusively, in full sun. 3-90 m

elevation.

Other material seen: PENINSULAR MALAYSIA. Johor: Kota Tinggi, 20 Jun 1934,

E.J.H. Corner SFN28612 (SING 0057465); Sungai Berassau, 8 Apr 1935, E.J.H.

Corner SFN29360 (SING 0031964; SING 0057453); Kota Tinggi, 9 Jul 1939, E./.H.

Corner SFN36965 (SING 0031943; SING 0057466); Kota Tinggi, 30 May 1937,

E.J.H Corner s.n. (SING 0057467); Pontian, 28 Sep 1939, 1. Ngadiman SFN36652 &

SFN36652a & SFN36652b (SING 0057468; SING 0057470; SING 0057471; SING

0031965; SING 0057469; SING 0057452); Kota Tinggi, 24 Jul 1939, 1. Ngadiman

(SING 0057462); Kota Tinggi, 28 Jul 1961, D.H. Nicolson 1223 (SING 0057475);

Johor, 1894, H.N.Ridley 6310 (SING 0057601); Tanjong Kupang, 1892, H.N.

Ridley s.n. (SING 0031913); Mt Austin, 1904, H.N. Ridley 12018 (SING 0031914);

Casthwood, H.N. Ridley s.n.(SING 0031963); Kukub, 1909, H.N. Ridley s.n. (SING

0057587); Johore, 17 May 1962, J. Sinclair 10690 (SING 0057463); Kota Tinggi-

- Mersing road, 2 May 1966, J. Sinclair 10895 (SING 0031944; SING 0057464).

Pahang: Sungai Lembing, | Jun 1964, H. Singh 1 (SING 0031946); Temerloh, 18 Mar

1923, M.R. Henderson 10527 (SING 0057449); Pahang, 6 Aug 1929, MR. Henderson

SFN22504 (SING 0057448); Pahang, 26 Jun 1891, H.N. Ridley s.n. (SING 0057450).

Melaka: Batang, Malaka, 2 May 1918 1H. Burkill 3193 (SING 0057461; SING

0057474; SING 0057476); Bukit Bruang, May 1901, C. Curtis s.n. (SING 0031862;

SING 0031863; SING 0031864); Melaka, 9 Apr 1891, PJ. Holmberg 743 (SING

0057472); Melaka, Aug 1892, H.N. Ridley 3529 (SING 0057473). Negeri Sembilan:

Jelebu, Pasoh F.R., 3 Jun 1987, J.V. LaFrankie LJV2271 (KEP); Tampin, 28 Jul 1915,

Mohd Nur 1317 (SING 0031945). Selangor: Kuala Langat, 4 May 1919, .H. Burkill

SFN4101 (SING 0057454); Telok F.R., 12 Jun 1921, 1H. Burkill SFN6521 (SING

0057459): Selangor, 10 Jun 1913, 4.M. Burn-Murdoch SFN155 (SING 0057447);

Selangor, Klang 3 Oct 1937, Mohd Nur SFN33992 (SING 0057455; SING 0057457;

SING 0057458); Telok F.R., 18 Jul 1961, D.H. Nicolson 1159 (SINGO031948); Telok

F.R., 18 Jul 1961, D.H. Nicolson 1160 (SING 0057456); Batang Berjuntai, May 1895,

H.N.Ridley 1662 (SING 0031947). Kuala Lumpur: Kuala Lumpur, 27 Jul 1889, HN.

Ridley s.n. (SING 0057460). Terengganu: Kuala Berang, 14 May 1925, R.E. Holttum

SFN15327 (SING 0057451). Perak: Batang Malacca, 1.H. Burkill SFN3192 (SING

0031860); Taiping, Larut, Jun 1882, H.H. Kunstler (Dr Kings collector) 3082 (KEP,

SING 0031910); Perak, 24 Jun 1961, D.H. Nicolson 1079 (K, SING 0057600, US);

Sungai Krian Estate, 21 Jun 1938, G.H. Spare SF 34592 (SING 0032091); Taiping, Jun

1888, L. Wray 2096 (SING 0031967). Penang: Province Wellesley, 15 Sep 1921, .H.

Burkill s.n. (SING 0057602).

SINGAPORE. Chua Chu Kang, 1905, H.N. Ridley s.n. (SING 0003668); Ang Mo Kio,

1894, H.N. Ridley s.n. (SING 0003671); Bukit Mandai, 1899, H.N. Ridley s.n. (SING

0003683); Singapore, 29 Jul 1934, Z. Teruya 2530 (KEP 38212).

BORNEO. SARAWAK: Kuching Division: Kuching, 19 May 1893 E./. Bartlett s.n.

(SING 0003758; 0003759); Bau, Kampung, Segong , 01°32°00.9"N 110°08°58.8”E,

10 Aug 2005, P.C. Boyce et al. AR-1333 (SAR); Lundu, Jalan Lundu, Kampung Perang,

01°37°18.9°N 109°53°05.7”E, 11 Sep 2005, P.-C. Boyce & Jipom ak Tisai AR-1355

(SAR); Lundu, Jalan Lundu, Stunggang Ulu 01°36’21.2°N 109°53°46.3”E, 21 Sep

2005, P.C. Boyce et al. AR-1372 (SAR); Lundu Sempadi, Gunung Papan, 19 Jun 2006,

———————————— ee

Helophytic Homalomena of Sunda

Lv)

LS)

P.C. Boyce & Jipom ak Tisai AR-1835, AR-1836 (SAR): Kuching, 15 Mar 1893, G.D.

Haviland 3134 (SING 0003767). Samarahan Division: Serian, Mongkos, Kampung

Kawan, 30 Nov 2005, P-C. Boyce & Simon Kutuh ak Paru AR-1535 (SAR). Sri Aman

Division: Sri Aman, Tempat Perkelahan Tapang Rumput, 23 Aug 2009 PC. Boyce

& Wong Sin Yeng AR-2593 (SAR). Mukah Division: Pulau Bruit, Sungai Kelepu,7

May 1957, J.A.R. Anderson 8012 (SING 0008587). Miri Division: Niah Suai, Niah

National Park, Trail to Great Cave, 03°49°21.7°N 113°45°44.7°E, 13 Oct 2005, PC.

Boyce, Jeland ak Kisai & Jipom ak Tisai AR-1416 (SAR). Limbang Division: Nanga

Medamit, Mulu N_P., trail from Kuala Likut to Camp 5, 04°07°02.3°N 114°49°26.9”E,

28 Sep 2007, P.-C. Boyce et al. AR-2223 (SAR); Nanga Medamit, Mulu N_P.. trail from

Camp 5 to Kuala Terikan, 04°12°58.0°N 114°53°20.1°E, 29 Sep 2007 PC. Boyce et

al. AR-2225 (SAR); Nanga Medamit, Mulu N.P. trail from Kuala Terikan to Camp 5

04°12°58.0°N 114°53°20.1”E, 3 Oct 2007 P.-C. Boyce et al. AR-2325 (SAR).

Etymology: Latin rostratus meaning beaked, curved, hooked, with a crooked point, in

referring to tip of the spathe limb prior to anthesis.

Notes on typifications: In publishing Homalomena sagittifolia Schott (1860) attributed

the trivial epithet to Junghuhn in schedula, implying that the name was appropriated

from a Junghuhn-annotated specimen in Bogor (“Java, Herb. Lugd. Bat.”). However,

Schott then went on to cite a Motley Borneo collection he had evidently seen as a

living plant presumably in Schoenbrunn (“v. v. — vide vivum™) and as a herbarium

specimen in Kew (“er s. in Herb. Hooker’). The Motley collection at Kew is fertile

but in poor condition, while the Junghuhn specimen appears to be no longer extant;

evidently the bulk of the description was prepared from the non-longer extant living

plant. The epitypification above is based on the only image in Schott’s Icones (W) to

bear both a Junghuhn and a Schott annotation of ‘Homalomena sagittifolia’ and is

most likely therefore to be the Junghuhn collection cited by Schott.

The two syntypes of Homalomena paludosa cited by Hooker (1893) are

incontrovertibly conspecific. The collection chosen here to serve as the lectotype is

in better condition, and has more distributed duplicates, and is thus the more suitable

choice.

Engler’s Homalomena raapii was published with two syntypes, of which only

the H.Raap collection cited above appears to be extant; it is assumed that the other

collections were destroyed during the bombing of Berlin herbarium during the Second

World War.

Alderwerelt’s Homalomena sagittifolia var. sumatrana is based on two

specimens, neither now in the best state of preservation. The specimen chosen is the

less degraded of the two.

Furtado’s Homalomena sagittifolia var. angustifolia is based on two syntypes.

Both extant. the collection chosen more closely matches the protologue and description,

and is in a better state of preservation.

Other notes: Homalomena rostrata is highly polymorphic in terms of leaf lamina shape,

and overall plant size, even within a single population, and has attracted a considerable

synonymy as a result. Flowering size plants range in height from barely 20 cm to over |

m, with the largest plants generally occurring in areas of abundant nutrient availability,

for example along the margins of freshwater swampforest, while smaller plants are

324 Gard. Bull. Singapore 62 (2) 2011

generally confined to nutrient-poor soils, especially on white sand and peat kerangas, or

along oligotrophic stream systems. Smaller plants tend to have sagittate leaves.

Leaf lamina morphology varies in two distinct manners: leaf width to length ratio,

with leaves ranging from sub-linear to oblong; and posterior lobe development, with the

leaf bases ranging from decurrent-cuneate through truncate to sagittate or hastate. These

morphologies are almost wholly mutually independent, and thus the base of a sub-linear

leaf lamina can range from decurrent-cuneate to hastate, while that of an oblong leaf

can range from truncate to sagittate or hastate. In extensive populations, e.g., at Niah, all

variations in plant size, and leaf lamina morphology, are represented.

Inflorescence morphology is far less variable, with any differences cited by

the authors of the synonyms readily encompassed by the variation in inflorescence

morphology acceptable in other far less narrowly defined species. In particular, abundance

or lack of flowers on the spadix and their relative disposition - congested or lax - and to

a great extent overall inflorescence size, is closely correlated to the size and vigour of the

plant producing the inflorescence.

Names now considered synonymous with H. rostrata can be assigned to the

following morphs:

— The robust, oblong-truncate leaf lamina morph includes the types of H. rostrata

Griff, H. miqueliana Schott, and H. miqueliana var. truella Alderw.

— The medium to robust, sub-linear leaf lamina morph includes the types of H.

beccariana Engl., H. ensiformis, and H. tevsmannii Engl.

— The robust oblong, sagittate to hastate leaf morph includes the types of H. sagittifolia

Jungh. ex Schott, H. sagittifolia var. pontederiifolia Ridl. (including the homotypic H.

ridlevana Engl.), H. sagittifolia var. angustifolia Furtado, and H. triangularis Alderw.

— The small growing, sagittate leaf morph is typified by H. propinqua Schott, H.

paludosa Hook.f., and H. sagittifolia var. sumatrana Alderw.

Helophytism in Araceae

Helophytism in Araceae is generally held to be a plesiomorphic condition, with

the greatest number of helophytic species occurring in subfamilies and tribes well

supported as being basal in the family: Orontioideae (three genera; eight species),

Lasioideae (seven of 10 genera; 26 of 36 species), Monsteroideae: Spathiphylleae

(one of two genera; many of the c. 60 species), and Calloideae (monospecific).

Helophytism in Aroideae is considered to be secondarily derived, with helophytic

taxa occurring scattered through most tribes: Spathicarpeae (Dieffenbachia Schott—

virtually all); Philodendreae (Philodendron Schott—several species, notably in

subgen. Meconostigma), Schismatoglottideae (arguably Phymatarum M.Hotta);

Cryptocoryneae (Cryptocoryne ciliata), Caladieae (Caladium Vent.—a few species);

Aglaonemateae (Ag/aodorum Schott—monospecific); Montrichardieae (all species);

Zantedeschieae (all species); Peltandreae (all species); Areae (7vphonium flagelliforme

(Lodd.) Blume); and Colocasieae (Colocasia Schott—a few, notably C. esculenta (L.)

Schott; A/ocasia (Schott) G.Don—a few, notably 4. a/ba Schott and A. sarawakensis

M.Hotta).

The occurrence of helophytism in Homalomena is thus unsurprising, although

it is interesting that the two species exhibiting this ecology are not considered

closely related. Homalomena expedita falls uncontroversially into the Homalomena

Helophytic Homalomena of Sunda 325

Supergroup (sensu Boyce & Wong 2008), while H. rostrata belongs in the Cyrtocladon

Supergroup (sensu Boyce & Wong 2008). Recently, chemical assaying has indicated

that the principal components of H. expedita are significantly different to those of

species in the Cyrtocladon Supergroup (Ng et al., in prep.): it is fully expected that

ongoing molecular work (Ng, et al., in prep.) will reinforce the recognition of two

distinct monophyletic lineages, and that helophytism is twice derived in Sundaic

Homalomena.

References

Boyce, P.C. & Wong, S.Y. (2008) Studies on Homalomeneae (Araceae) of Borneo I:

Four new species and speculation on informal species group in Sarawak. Gard.

Bull. Singapore 60(1): 1-29.

Boyce, P.C. & Wong, S.Y. (2009) Studies on Homalomeneae (Araceae) of Borneo

IV: Homalomena specimens in the Herbarium Beccarianum-Malesia (FI-B) of

the Museo di Storia Naturale - Sezione Botanica “F. Parlatore™ dell’ Universita di

Firenze. Webbia 64(2): 169-173.

Boyce, P.C., Wong, S.Y. & Fasihuddin, B.A. (2010) Studies on Homalomeneae

(Araceae) of Borneo II: The Homalomena of Nanga Sumpa (Batang Ai) — Novel &

pre-existing taxa, and notes on Iban Usages. Gard. Bull. Singapore 69(2): 29-78.

Hay, A. & Herscovitch, C. (2002) Two Remarkable New West Malesian Homalomena

(Araceae) Species. Gard. Bull. Singapore 54(2): 171-178.

Hooker, J.D. (1893) Araceae. In: J.D. Hooker. The Flora of British India 6: 490-556.

London: Reeve & Co.

Schott, H.W. (1860) Prodromus systematis Aroidearum. Vienna: Typis Congregationis

Mechitharisticae.

Gardens’ Bulletin Singapore 62 (2): 327-328. 2011 B27]

Ginalloa siamica var. scortechinii is a

species of Viscum (Viscaceae)

HE Yao

Forest Research Institute Malaysia, 52109 Kepong, Selangor, Malaysia

yaotzeleong@frim.gov.my

ABSTRACT. Ginalloa Korth. is not represented in Peninsular Malaysia. Ginalloa siamica

Craib var. scortechinii Gamble, known only from the type specimen, is conspecific with Viscum

ovalifolium Wall. ex DC.

Keywords. Ginalloa, Peninsular Malaysia, Viscaceae, Viscum

Introduction

In Peninsular Malaysia, Ginalloa Korth. is recorded only through Ginalloa siamica

Craib var. scortechinii Gamble (Gamble 1914) described from a single specimen,

Scortechini s.n., Perak (CAL, accession number 396346). According to Barlow

(1997), Ginalloa siamica occurs in Thailand, Cambodia, Peninsular Malaysia and the

Philippines. Variety scortechinii is distinguished by its 3-veined leaves compared to

the 5-veined ones in the typical variety.

Notes on Ginalloa siamica var. scortechinii

The Scortechini type specimen is undated and the exact locality not specified.

According to Gamble (1914), Ginalloa is characterised by flowers in triads on terminal

and/or axillary spikes and female flowers with 3 minute perianth lobes, contrasting

with Viscum L. that has flowers which are usually in triads or sometimes solitary, in

the axils of leaves or at leafless nodes on branches, and female flowers usually with

4 perianth lobes. As the specimen Gamble described has a spike-like inflorescence

bearing only female flowers with 3-lobed perianths, he described it as Ginalloa.

However, Gamble’s note (19 Nov 1912) on the specimen reads: “This is the

only sheet available. I found it among the sheets of Viscum orientale. Other sheets

should be searched for and some sent to the Kew Herbarium.”

I searched the Peninsular Malaysian Viscum collection in the Kew Herbarium

and among specimens of Viscum ovalifolium Wall. ex DC. (formerly Viscum orientale

auct. non Willd.) that have 3-veined leaves, I found a collection (Scortechini 732)

identical to the type of Ginalloa siamica var. scortechinii. Detailed examination

showed that the shoot branching, leaf size, shape and venation, and fruits in leaf axils

and at leafless nodes were the same. I therefore conclude that Ginalloa siamica var.

scortechinii is synonymous with Viscum ovalifolium and Ginalloa does not occur in

Peninsular Malaysia.

328 Gard. Bull. Singapore 62 (2) 2011

Barlow (1997) noted that Viscum ovalifolium is polymorphic, and that while its

flowers are usually arranged in triads or sometimes solitary, they are less usually found

along leafless shoots that superficially resemble the spicate inflorescences of Ginalloa.

This is well illustrated by the collection Chew et al. FRI 53735 (KEP). Also, it is not

uncommon for female flowers to have a perianth lobe missing, i.e. with only three

lobes present. Danser (1931) had maintained Ginalloa siamica var. scortechinii but

Barlow (1997) did not recognise the variety, although the latter still regarded Ginalloa

siamica as having a distribution that extended into Peninsular Malaysia. Neither had

seen the type (Barlow, pers. com.).

Viscum ovalifolium Wall. ex DC., Prodr. (DC.) 4: 278 (1830). TYPE: Wallich 489,

Penang (holotype K-W).

Ginalloa siamica Craib var. scortechinii Gamble, J. Asiat. Soc. Bengal 75: 383

(1914), syn. nov.

_ TYPE: Scortechini s.n., Perak, Peninsular Malaysia (holotype CAL, accession number

396346).

ACKNOWLEDGEMENTS. I am deeply indebted to M.S. Mondal from the Botanical Survey

of India, Central National Herbarium (CAL) for providing the image of the specimen. I am

grateful to the curators of the herbaria of the Royal Botanic Gardens, Kew (K), Nederlands

Centre for Biodiversity Naturalis (section Nationaal Herbarium Nederland), Leiden University

(L), and Singapore Botanic Gardens (SING) for permission to examine specimens in their care.

The Ministry of Science, Technology and Innovation of Malaysia is acknowledged for financial

support for the Flora of Peninsular Malaysia Project (No. 01-04-01-000 Khas 2) under which

this study was carried out. I am grateful to L.G. Saw, R.C.K. Chung and R. Kiew for advice and

comments in improving the manuscript. Also, I would like to express my gratitude to one of the

reviewers who took time to do a careful editing in helping me to shorten the text.

References

Barlow, B.A. (1997) Viscaceae. F7. Malesiana, Ser. I, 13: 403-442.

Candolle, A.P. De (1830) Prodromus systematis naturalis regni vegetabilis 4. Paris:

Treuttel & Wiirtz.

Danser, B.H. (1931) The Loranthaceae of the Netherlands Indies. Bull. Jard. Bot.

Buitenzorg IIT, 11: 233-519.

Gamble, J.S. (1914) Loranthaceae. Flora of the Malayan Peninsula. J. Asiat. Soc.

Bengal 75: 349-391.

Gardens’ Bulletin Singapore 62 (2): 329-330. 2011 329

BOOK REVIEW. Trees of Tropical Asia. An Illustrated Guide to Diversity. James

V. LaFrankie, Jr. 2010.

Philippines: Black Tree Publishing, Inc. 22.9 cm x 15.5 cm, card cover. 750 p. ISBN 978-97 1-—

94794—0-6. Price SGD 125 / US$ 149.

The author has 20 years of field experience in several countries of SE Asia, including

a considerable time in the company of the legendary Malaysian botanist K.M.

Kochummen. This collaboration must have been very profitable for the author, which

probably explains why Kochummen is the only person consistently referred to as

euir:”’.

SE Asia is defined as the tropical area from Burma to Borneo and the Philippines.

The flora in this part of the world is so rich that a choice has to be made: 157

families and 887 genera are treated, with notes on description, ecology and possible

confusion. Numerous illustrations, mostly of good quality, help to give an idea of the

taxa discussed. Some genera not occurring in “SE Asia” are also included, such as

Schumacheria, a Sri Lankan endemic; Eriandra, a Papuasian endemic; Galbulimima;

Pigafetta and Macadamia, extending west to Sulawesi.

Although the book deals principally with trees, also lianas, shrubs and even

some herbs are mentioned, giving the reader more than promised. On the other hand

I wonder why some prominent woody taxa such as Dracaena and the omnipresent

Bambusoideae are left out.

The book is said to be aimed at beginning students, young staff of local herbaria

and overseas botanists who want to know more about Asian plants. To this aim the

author has condensed a large amount of information by using a very small lettertype

and leaving out author names. In the introductory chapters the author explains that

modern molecular and cladistic research has led to changes in traditional systematic

concepts. Although the new phylogeny often confirms some “suspicions” based on

morphological grounds, more often than not the conclusions can be quite baffling.

Who would suspect that Rafflesia is closer to Euphorbiaceae s.s. than Daphniphyllum

or Dichapetalum? On the other hand, the author allows morphological evidence to

prevail in the case of Bischofia. This genus belongs to the Phyllanthaceae (split off

from Euphorbiaceae) on molecular grounds, but is placed in a separate family because

it is morphologically aberrant.

The author urges his readers to look at trees as living populations and not

as dried specimens in the herbarium. He also warns that DNA-based information,

albeit necessary and sensible, should not lead to the assumption that inventory and

enumerations are in the eclipse. I would like to add that young botanists are perhaps

best advised to learn to recognise genera rather than families. The last word in modern

family phylogeny is far from being said. Generic concepts seem to be less prone to

changes.

An extensive list of references is given in which, to my surprise, many

prolific authors of the Flora Malesiana series (Ding Hou, Leenhouts, Sleumer) are

not mentioned. Indices to scientific names are added as well as to Malay, Thai and

Vienamese names, but not to Indonesian names. The few Indonesian names in the text

are often wrong. My main point of criticism is the number of spelling and printing

errors.

330 Gard. Bull. Singapore 62 (2) 2011

All the same, I want to compliment Dr. LaFrankie for bringing together a vast

amount of information on one of the most diverse floras of the world. It is the first

work of its kind.

Max van Balgooy

Nationaal Herbarium Nederland

Gardens’ Bulletin Singapore 62 (2): 331-332. 2011 331

BOOK REVIEW. Flora of Peninsular Malaysia. Series II: Seed Plants, Volume 1.

(Malayan Forest Records No. 49) R. Kiew, R.C.K. Chung, L.G. Saw, E. Soepadmo &

P.-C. Boyce (eds). 2010.

Kepong: Forest Research Institute Malaysia. 25.7 cm = 18 cm, hard cover. 329 p. ISBN 978-—

967-5221—32-3. Price RM 100 / USS 75.

This is the first volume of the long-awaited Flora in the wake of the Tree Flora of

Malaya and Tree Flora of Sabah and Sarawak. The volume is well structured,

beginning with a historical overview of botanical interest in the area, followed by

highlighting the latest family realignments adopted and the families affected.

Subsequent to that is an account of the vegetation of Peninsular Malaysia and current

conservation measures, as well as criteria used to categorise the conservation status of

species, before embarking on the Flora proper.

‘A Brief History of Taxonomic Research in Peninsular Malaysia’ (Kiew. Chung,

Saw & Soepadmo) provides an insight into the botanical interest, collecting activity,

as well as the establishment of various herbaria in old Malaya (today’s Peninsular

Malaysia and Singapore), chronologically listing many botanists who have done

taxonomic research work pertinent to this region, from Hooker to the present day

and indicating why a flora account is long overdue since Ridley’s five-volume work,

The Flora of the Malay Peninsula. In the account that highlights taxonomic changes

in family composition, as well as the recognition of families based on molecular

phylogenetic work, the inclusion of a phylogenetic tree showing the evolutionary

relationships of the affected families could have been helpful. Some useful statistics,

such as the number of species of seed plants in Peninsular Malaysia (estimated 7,834

species with 1,564 genera in 220 families), with detailed breakdown, are provided

with the families arranged alphabetically, such that the number of species and genera

in the families at a glance are available (pages 16-20). The account of vegetation

(Saw), well compiled and easy to follow, is a useful accompaniment to forming better

impressions of the ecology of the species in this flora. L.S.L. Chua gives a brief outline

of species assessment and conservation in Peninsular Malaysia.

The format of the enumeration comprehensively provides accepted name,

etymology, key references and type citation, description, vernacular name and

distribution. Identification keys are well-formed and provided when applicable, but

it is sometimes clear that it has been necessary to depend on reproductive instead of

vegetative characters. Mention of the uses of the species, a species distribution map,

conservation status, ecology, taxonomic notes and the italicising of key characters make

the account informative and concise. Given under ‘Notes’ are nuggets of information

either to provide some identification tips, clarify misconceptions or provide further

useful information pertaining to a species, such as phenological attributes.

In this first volume, the families treated are: Ancistrocladaceae, Araucariaceae,

Balanophoraceae, Bonnetiaceae, Casuarinaceae, Chloranthaceae, Clethraceae,

Cruciferae (naturalised species only), Ctenolophonaceae, Daphniphyllaceae,

Datiscaceae, Erythroxylaceae, IIliciaceae, Myricaceae, Nelumbonaceae, Pedaliaceae

(naturalised), Pentaphylacaceae, Pittosporaceae, Podocarpaceae, Portulacaceae (many

escapes and weeds), Schisandraceae, Symplocaceae, Tetrameristaceae, Torricelliaceae,

Trigoniaceae, and Turneraceae (naturalised).

Overall, this may be considered a well-illustrated volume. On the first page,

there is a useful reference coloured map of Peninsular Malaysia indicating general

332 Gard. Bull. Singapore 62 (2) 2011

Sap

topography and drainage. The demarcations of the states are not well defined in certain

areas, though. Another map on page 25 that shows the vegetation should be enlarged

to full-page, as the text is too small for comfortable reading. Included are 17 pages of

colour plates and 42 well-drawn figures of species treated, in varying detail.

This flora is long overdue, not just useful to the taxonomist, but also others

interested in identifying plants. It is accessible also to students and others wishing to

know more about the rich plant life of Peninsular Malaysia.

Paul K.F. Leong

K.M. Wong

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 62 (2): 333-334. 2011

1S)

BOOK REVIEW. Malaysia Plant Red List, Peninsular Malaysian Dipterocarpaceae.

(FRIM Research Pamphlet No. 129) L.S.L. Chua, M. Suhaida, M. Hamidah & L.G.

Saw. 2010.

Kepong: Forest Research Institute Malaysia. 25.5 cm = 18.5 cm, card cover. 210 p. ISBN

978—967—-5221—34-7. Price RM 20.00 / USD 15.00.

In view of the worldwide threat to our environment through habitat degradation and

land use change, conservation efforts with the aim to protect the world’s biodiversity

have increased dramatically over the last few decades. They are generally based on

the understanding that any measures to ensure successful conservation and sustainable

utilisation of the biclogical diversity require a detailed understanding of the plant and

animal species to be protected, including their distribution, ecology and conservation

status. In Malaysia, one of the aims of the National Strategy for Plant Conservation

is to obtain preliminary assessments of the conservation status of all plant species of

the country, and as a consequence a project on the conservation monitoring of rare

and threatened plants was initiated. Families Dipterocarpaceae, Palmae (= Arecaceae)

and Begoniaceae have been identified for the first phase of this project. Being an

economically important ‘keystone family’, the dipterocarp family is treated in the

present book which is the first in a series of Red Data assessments.

Although many Malaysian tree families are included in a recent global IUCN

Red List, there is currently no national Red Data List, and the present publication

is intended to fill this gap for the dipterocarp family. The introductory part of this

book gives background information on family Dipterocarpaceae (distribution, habitat

ecology, uses), on land use changes in Malaysia and threats to dipterocarp trees. A

section explains the IUCN Red List Categories and Criteria version 3.1 (2001) that

have been used in the assessments. Existing conservation policies and laws are

discussed, and future steps in the conservation monitoring are outlined. The scope

and objectives of the conservation monitoring project which forms the basis for

the present book are explained. The present Red List is intended as the first critical

step towards prioritisation of national conservation measures. Several conservation

recommendations are made, most importantly the need to incorporate conservation

principles at the planning stage of developments, to make additions to legal and policy

provisions with regard to habitat protection, to further fundamental research to increase

our knowledge of threatened species, to organise educational programmes aimed at

local involvement, and to increase networking efforts among stakeholders. One of

the important outcomes of the conservation monitoring project is that the cooperation

between the Forest Research Institute Malaysia (FRIM) and various stakeholders

has resulted in an increased awareness of rare and threatened species and the need

to conserve them. Rescue operations are encouraged which, apart from rescuing

threatened populations, would also increase ex situ collections of Dipterocarpaceae.

Three informative tables are given in the introductory part, showing the number

of taxa arranged after genera in each of the IUCN categories, listing all of the threatened

taxa, and breaking down the number of threatened taxa after the Malaysian states.

Ninety-two taxa occurring in Peninsular Malaysia are considered threatened, which

is roughly 56% of the total number, while one (Shorea kuantanensis) is considered

extinct. The Malaysian state with the largest number of threatened taxa is Pahang with

59, followed by Johor (54) and Perak (53). Compared with previous assessments, the

conservation status of six taxa has been upgraded to a higher threatened category, and

334 Gard. Bull. Singapore 62 (2) 2011

in 57 taxa downgraded to a lower threatened category. Fifty-eight taxa which were

assessed as threatened before were here downgraded to not threatened. Another result

of the present conservation monitoring project is that one new species was discovered

(Vatica yeechongii) and two new distribution records were made (Dipterocarpus

tempehes, Dryobalanops beccarii). Nine species which were previously thought to be

extinct were rediscovered.

In the main section of the book, the conservation status of all except one of

the 165 taxa of Dipterocarpaceae occurring in Peninsular Malaysia is assessed

according to the IUCN Red List Categories DD (= Data Deficient), EX (= Extinct),

CR (= Critically Endangered), EN (= Endangered), VU (= Vulnerable), NT (= Near

Threatened) and LC (= Least Concern). In the three threatened categories CR, EN

and VU the assessment criteria are indicated, e.g., VU B2b(ii1)+c(11). The assessments

are based on information obtained from specimens found in Malaysian as well as

in European herbaria and on fieldwork during the present conservation monitoring

project. In addition, further contributions have been made through data obtained in

- decades of forest management. Taxa that occur in both Peninsular Malaysia and East

Malaysia (= Sabah + Sarawak) are assessed separately for each region, and the higher

category was chosen for the national conservation status. In each of the 164 assessed

taxa the vernacular names, distribution (both worldwide and within Malaysia) and

habitat are also given. Various additional conservation notes are also provided (mostly

referring to the presence of the taxon in protected areas), and previous assessments are

cited. Each one of the assessments is accompanied by distribution maps of Peninsular

Malaysia and East Malaysia (1f occurring here also). Grey shading in these maps

indicates, a) in case of Peninsular Malaysia the extent of the forest cover of the region,

and b) in case of East Malaysia the Extent of Occurrence of the dipterocarp taxon

(using grey shading for different attributes in the maps seems somewhat confusing

to me, but this is perhaps just a matter of personal preference; in any case it is the

only negative point noted in this review). Attached to these maps are several selected

localities, the Extent of Occurrence of the taxon, the Area of Occupancy of the taxon

and, for Peninsular Malaysia, also the extent of the forest cover within the Extent of

Occurrence. In many of the distribution maps also ex sitw collections are marked. For

a number of taxa, line drawings of various plant features (leaves, flowers, fruits) are

provided in this assessment section.

The text is followed by a large section with high-quality colour illustrations,

comprising well over 150 photos grouped in 39 plates. A total of 68 taxa are illustrated.

Most of the photos show the bark, leaves and young or mature fruits, and in some cases

also the entire tree, the flowers or seedlings are shown. An index to scientific names

and another to vernacular names conclude the book.

The book is the first complete assessment of the conservation status of

Peninsular Malaysian dipterocarps, and, with its clear and well-arranged text and the

many informative distribution maps and plant illustrations, should be recommended to

everyone with an interest in tropical trees.

Hubert Kurzweil

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 62 (2): 335-336. 2011 335

BOOK REVIEW. Alexander von Humboldt and the Botanical Exploration of the

Americas. H. Walter Lack (translated from the German by Stephen Telfer, Edinburgh).

2009.

London: Prestel Publishing Ltd. 278 p. ISBN 978-3-7913-4142-2. Price £ 125.

This book is, quite simply, a sumptuous treat to indulge oneself. Beautifully produced

on quality paper, boxed in a fine slipcase, the colour reproductions a delight to the

eye, it has been a long time since I so enjoyed the sensory experience of exploring a

new book such as this. And it is a compliment to the publishers. Prestel Verlag and

their international offices, that such beautiful books are still being produced, albeit at

a price.

So who is Alexander von Humboldt, the subject of this beautiful tome, and why

_would a reader in Asia care about his explorations of the Americas? To those who are

not familiar with the name, Humboldt (1769-1859) was one of the greatest explorers of

the early nineteenth century, a man who pioneered the field methods now used around

the world in disciplines as diverse as botany, geology, geography, and oceanography.

He belonged to a generation of European scientists that was no longer content to

remain cocooned in an academic institute lecturing about places he had never seen

on subjects he knew only from second hand. Instead he traveled to tropical America

and endured 5 years of hardship to see first-hand the lands, peoples, cultures, and

biodiversity there. Humboldt’s name is commemorated in a multitude of geographic

features and living things; indeed, virtually any landform, oceanic phenomenon, plant,

or animal that includes ‘Humboldt’ is surely named in honor of this German scientist

and philosopher.

The author of this book, Professor H. Walter Lack, is the pre-eminent scholar

of Humboldt’s botanical legacy, but until now his publications have been entirely in

German. Prestel’s decision to translate Lack’s book into the English language makes

the story of the expedition and the importance of its scientific contributions more

widely available.

The book begins with a brief overview of Humboldt’s early life and education,

then takes up the story of the “great expedition” to the Americas. This lasted for five

years (1799-1804) and is summarised succinctly in 18 pages that include numerous

colour illustrations. The rationale for this brief treatment is that, in the author’s words,

“The story of the expedition ...has been told ad nauseam...” While this is borne out via

the references cited in the German language. an opportunity has been missed here to

describe the expedition more fully to readers in the English language. Nonetheless,

the synopsis presented hits the high points and gives a very cogent explanation of

the scientific field methods employed by Humboldt and his companion and botanical

collaborator, Aimé Bonpland, methods that are largely still followed today by botanists

collecting plants in the field.

Then follow seven chapters dealing in some detail with the scientific publications

that followed after the expedition, a chapter covering the later years of Humboldt and

his botanical collaborators Bonpland and Carl Sigismund Kunth. a summary of the

precise dates of publication for the various works mentioned in these chapters, and a

postscript explaining the big picture of what the publications accomplished and musing

on the roles of the various principles. While these might seem dry and uninteresting,

the reality is that they make for fascinating reading. The complex personalities and big

egos, political tides in Europe. and nationalistic tussling assured that the publication

336 Gard. Bull. Singapore 62 (2) 2011

of results was anything but simple and the scientific outcomes are still resonating in

taxonomy today. Indeed, those who enjoy a good detective novel will find the story of

the post-expedition publications intriguing.

For those more visually inclined, the next portion of the book will be the

focus of interest: 82 magnificent plates reproduced from the botanical publications

that came out of the collaborations and competitions. Some of these are black and

white engravings, others are full-colour plates; all are superbly reproduced and are, no

doubt, the principle reason for the cost of this book. Lovers of art and plants will find

them fascinating; most are credited to the botanical artist Pierre Turpin, a few are the

work of Humboldt himself (see plate 33, an orchid, for example)—he was trained as

a technical draughtsman and illustrator among his many other accomplishments—and

the remainder are the work of noteworthy botanical artists including Redouté, Delile,

and others.

One error was detected: in the caption for plate 42 there is a transposition of

letters in the name of the genus between the plate and the caption; Aragoa on the

- plate, which is correct, became Aragao in the caption. This lapse aside, the plates are

beautifully reproduced and give a glimpse of the botanical riches the great expedition

gathered and described.

The book concludes with a bibliography of selected works about Humboldt and

a series of three indexes, to places, proper names, and botanical names for plants. My

compliments to the author and the publisher: this book is a welcome and beautiful

addition to the body of work about Alexander von Humboldt and his botanical

contributions to science.

George Staples

Singapore Botanic Gardens

Gardens’ Bulletin Singapore 62 (2): 337-338. 2011

(oS)

Le)

BOOK REVIEW. Bamboos at TBGRI. K.C. Koshy. 2010.

Thiruvananthapuram. Kerala, India: Tropical Botanic Garden and Research Institute. 27.5 cm

x 19.8 cm, card cover. 104 p. Price Rs. 800 / USS 30.

“All botanic gardens keep records of their living collections.” Thus declares the opening

sentence in the preface to this book by Dr. K.C. Koshy, the founder of the Bambusetum

at the Tropical Botanic Garden and Research Institute (TBGRI) in Kerala, India. That

exhortation does not detail how well those records should be kept, and Koshy proceeds

to show how.

This account was well-conceived, at a time when the TBGRI bamboo collection

has just entered its third decade with nearly 70 species (currently recognised as 15

genera) and more than ten putative hybrids making up a whopping 933 accessions.

a very good stage at which to declare coming of age. The opening chapter sets the

foundation of the Bambusetum in the context of TBGRI’s history. Beginning with

a mere 0.5 ha at inception, this Bambusetum has now grown to occupy nearly 6.6

ha (just more than 16 acres) in the sprawling green 121-ha campus of the TBGRI.

Koshy gives a quick overview of Asian and Indian bambuseta, drawing attention to

the humble beginnings of those collections that today continue to support scientific

studies, such as at the East India Company (later Indian) Botanic Garden at Howrah,

Calcutta from the time of Roxburgh in the early 19th century. as well as a good

number of newer collections established around the 1980s. Notes are provided on

the propagation methods used at TBGRI, the stage-by-stage development. and the

geographical provenances covered that emphasised the Western Ghats region and

North Western and North Eastern India. Koshy also documents how thirteen taxa

were duplicated from the bambusetum of the Forest Research Institute at Dehra Dun,

where specialist attention for bamboo taxonomy and genetics, biology and utilisation,

have been and continue to be emphasised. This kind of duplication represents some

insurance against loss at one location and also allows comparative studies in different

environments. One or two anecdotes, such as the difficulties of transporting bulky

plant offsets by passenger train, the Manipur collecting expedition through insurgency

areas, or the monsoonal damage to Bambusetum plants in June 2003 that had to be

dealt with, make the account come alive. Building a bambusetum is not just planting

bamboo, it brings us into contact with much else, and often in ‘heavyweight’ fashion.

The main records are compiled as Chapter 2. Herbarium and spirit-preserved

material help document the living accessions. The entries are organised by genus and

species in alphabetical order. Each taxon entry provides the scientific name, relevant

taxonomic references, a brief description of the species, a literature-based distribution

statement, and a note on the number of hereditary lines represented by the TBGRI

material. The accession numbers of the taxon are given with location statement and

(presumably GPS) coordinates in the TBGRI Bambusetum: planting date; propagule

type used: details of origin (typically state, district and precise location), collection

date, and collector and collection number. Herbarium and spirit collections associated

with the material (either collected from outside the Bambusetum, or within: this is

carefully distinguished) are also given when available. But not only are there planted

bamboo accessions in the Bambusetum, as two natural populations, Bambusa bambos

and Ochlandra travancorica, have been carefully conserved on site.

This book is very nicely produced, replete with end-paper photographic spreads

of Pseudoxytenanthera bourdillonii (Gamble) H.B.Naithani and a frontispiece with

338 Gard. Bull. Singapore 62 (2) 2011

an alluring teaser in the form of an unidentified gigantic Dendrocalamus sp. Nearly

every page has full-colour photographs, and there are a good number of full-page

photographic reproductions (a Roxburgh painting of Melocanna baccifera faces the

Preface), including many of the Bambusetum accessions. A contour map of the TBGRI

Bambusetum showing the locations of all accessions spans nearly two pages, which

means a serious fold is found right across it, and the fine print used for accession

numbers on this map can be a challenge to the reader. Apart from this, the book is a

highly unusual, but extremely welcome, detailed (and pleasant-to-consult) record of a

scientific collection of living bamboos. It is unprecedented.

The TBGRI is a young institution, set up in 1979, and its Bambusetum was

established in 1987, so it could be surprising that, in effect, this book unveils to the

modern world one of the finest, if not the best, scientific living collections of bamboo

in all of India and the tropics. And K.C. Koshy, certainly, has set the standard for not

only TBGRI, which looks towards an ever-increasing role in tropical plant research,

but any institute that seriously wishes to build a scientific collection of bamboos.

TBGRI has had the ‘right recipe’: forefathers with good breadth of vision, passionate

researchers who are well-qualified and dedicated to the task, good networking, and the

great Kerala setting for tropical plants. It had worked before in the tropics and works

again.

K.M. Wong

Singapore Botanic Gardens

Instructions for contributing authors (continued)

Title and authorship. The title should concisely describe the contents. If a scientific name is used, its

authority is normally excluded, but the family name would be provided. Authors’ names, affiliations

and postal / e-mail addresses are stated below the title. If more than one author, indicate “corresponding

author”. Avoid footnotes. A short running title (up to six words) should also be provided.

Abstract. The abstract is at most 100-300 words. It should concisely indicate the article’s contents without

summarising it; mentioning novelties and name changes. Keywords: Suggest at most eight keywords, in

alphabetical! order.

Scientific names and author abbreviations. Genus and species names of organisms must be italicised

and followed by the authority (with family name in parentheses) when first mentioned in the text or

diagnoses. Standards for author abbreviations include:

Brummitt, R.K. & Powell, C.E. (1992) Authors of Plant Names. Kew: Royal Botanic Gardens, Kew.

Kirk, P.M. & Ansell, A.E. (1992) Authors of Fungal Names [Index to Fungi Supplement]. Wallingford:

CAB International.

Herbarium abbreviations (http://sciweb.nybg.org/science2/IndexHerbariorum.asp) follow

Holmgren, P.K., Holmgren, N.H. & Barnett, L.C. (eds) (1990) Index Herbariorum. Part I: The Herbaria

of the World [Regnum Veg. vol. 120]. New York: New York Botanical Garden.

Journal and book title abbreviations. For journals:

Bridson, G.D.R., Townsend, S.T., Polen, E.A. & Smith, E.R. (eds) (2004) BPH-2: periodicals with

botanical content: comprising a second edition of Botanico-Periodicum-Huntianum. Pittsburg: Hunt

Institute for Botanical Documentation.

For books:

Stafleu, F.A. & Cowan, R.S. (eds) (1976-88) Taxonomic Literature. 2nd edition. 7 vols [Regnum Veg.

vols 94, 98, 105, 110, 112, 115, 116].

Stafleu, F.A. & Mennega, E.A. (eds) (1992—) Taxonomic Literature. Supplements [Regnum Veg. vols

125, 130, 132].

(A useful source of verifying names of publications is

<http://asaweb. huh. harvard.edu:8080/databases/publication_index.html>. If in doubt, list full titles.

Other abbreviations and units of measurement. If using standard abbreviations and acronyms, give the

full term on first mention. Dates are cited as: | Jan 2000. SI (metric) units of measurement are used and

spelled out except when preceded by a numeral; they are abbreviated in standard form: g, ml, km, etc.

Tables. Tables are numbered in arabic numerals in the order they are first mentioned in the text and carry

an indicative legend at the head. Tables are given at the end of the manuscript.

Illustrations. All drawings, maps, graphs and photographic images (individually or collected in a plate)

are to be numbered in arabic numerals in the order they are first mentioned in the text, as Fig. 1, Fig. 2,

etc. (plate components would be referred to in the text as Fig. 1A, 1B, Fig. 1A—D, etc.). If relevant, scale

bars should be used to indicate magnification.

When grouping photographs, the maximum page area 19.5 x 13 cm must be heeded. High resolution digital

images may be submitted as separate files (line drawings in black and white at 600 dpi, photographs at

300 dpi) sent electronically or in a CD. Do not embed images into the main text file.

References in the text. Citation in the text should take the form: King & Gamble (1886) or (King &

Gamble 1886), or King et al. (1886) if more than three authors to a work. Use 2000a, 2000b, ete. if

several papers by the same author(s) in one year are cited.

References listed at the end. There, works mentioned in the text are listed alphabetically as follows:

Dallwitz, M.J., Paine, T.A. & Zurcher, E.J. (1999) User’s Guide to the DELTA Editor. http:/

biodiversity.uno.edu/delta/ (accessed on 2 Aug. 2010).

Persson, C. (2000) Phylogeny of Gardenieae (Rubiaceae) based on chloroplast DNA sequences from the

rps\6 intron and trnL(UAA)-F(GAA) intergenic spacer. Nordic J. Bot. 20: 257-269.

Ridley. H.N. (1930) The Dispersal of Plants Throughout the World. Ashford, U.K.: L. Reeve.

Smith, A.C. & Darwin, S.P. (1988) Rubiaceae. In: Smith, A.C. (ed) Flora Vitiensis Nova, A New Flora

of Fiji 4: 143-193.

References to web-based resources should include either a doi (digital object identifier) specification

or full URL mentioning also the date it was accessed. Use of DNA sequences from GenBank should be

acknowledged and the studies for which the sequences were generated should be cited.

Style of nomenclatural summaries. The following style is required:

Gardenia anisophylla Jack ex Roxb., Fl. Ind. ed. Carey & Wall. 2: 561 (1824).

Medinilla alternifolia Blume, Mus. Bot. 1: 19 (1849).

If authors include full bibliographic data for these works in the list of references at the end of the paper,

they should also be mentioned in the text briefly, e.g., “Nomenclatural references researched include

Blume (1849) and Roxburgh (1824).”

ee Saunas should be ee in a block, stating the type at the end.

Front cover picture: Medinilla tapete-magicum Camara-Leret & Veldk. (Photo al Irawat1)