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T) Singapore
Botanic Gardens
he Gardens^’ Bulletin
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VOL. 66(2) 2014
7859
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THE GARDENS’ BULLETIN, SINGAPORE
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conservation, with emphasis on the plant life of the Southeast Asian-Pacific region.
Dr David J. Middleton Dr Jana Leong-Skomickova
(Editor-in-Chief) (Managing Editor)
S. Lee Y.W. Low Christina Soh
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Editorial Advisory Board
Professor Sir Peter Crane
Yale University!
USA.
Dr Rogier P.J. de Kok
Royal Botanic Gardens, Kew
UK.
Dr W. John Kress
National Museum of Natural History
Smithsonian Institution
USA
Dr Mark Hughes
Royal Botanic Garden Edinburgh
UK.
Dr Kiat W. Tan
Gardens By The Bay
Singapore
Dr Nigel P. Taylor
Singapore Botanic Gardens
National Parks Board, Singapore
Dr Ian M. Turner
Singapore Botanic Gardens
National Parks Board, Singapore
Dr Jan-Frits Veldkamp
National Biodiversity Center
The Netherlands
Dr Jun Wen
National Museum of Natural History
Smithsonian Institution
US.A.
Professor Nianhe Xia
South China Institute of Botany
PR. China
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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 16 intron and ?mL(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. & Damin, S.P. (1988). Rubiaceae. In: Smith, A.C. (ed) Flora Vitiensis Nova, A New Flora
of Fiji A. 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:
Ornithoboea arachnoidea (Diels) Craib, Notes Roy. Bot. Gard. Edinburgh 11: 251 (1920); Burtt, Notes
Roy. Bot. Gard. Edinburgh 22: 294 (1958).
Ornithoboea parishii C.B. Clarke in A. DC. & C.DC., Monogr. Phan. 5(1): 148 (1883).
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 tlie text briefly, e.g., “Nomenclatural references researched include
Blume (1849) and Roxburgh (1 824).”
Homotypic synonyms should be provided in a block, stating the type at the end.
Front cover picture: Zingiber singapurense (Photo by Jana Leong-Skomickova)
The Gardens’ Bulletin
Singapore
VOL. 66(2) 2014
ISSN 0374-7859
CONTENTS
Y.R Deng
Phlogacanthus magnus, a new combination in the Acanthaceae
from Malaysia 121
R. Kiew
Two new species and one new subspecies of Ridleyandra (Gesneriaceae)
from Peninsular Malaysia 125
R. Kiew
Two new white-flowered Codonoboea species (Gesneriaceae)
from Peninsular Malaysia 137
J. Leong-Skornickova
Anew lectotypification of Zingiber gracile var. elatius (Zingiberaceae) 145
J. Leong-Skornickova, A. Thame & P.T. Chew
Notes on Singapore native Zingiberales I: Anew species of Zingiber
and notes on the identities of two further Zingiber taxa 153
S. Lindsay 4& C.W. Chen
Three new combinations in Haplopteris (Pteridaceae subfam. Vittarioideae) 169
R.O. Makinson & B.J. Conn
Puccinia psidii (Pucciniaceae - Eucalyptus Rust, Guava Rust, Myrtle Rust)
- a threat to biodiversity in the Indo-Pacific region 173
D.J. Middleton, J. Leong-Skornickova & Q.B. Nguyen
A new species of Billolivia (Gesneriaceae) from Vietnam 189
M. MoUer, W.-H. Chen, Y.-M. Shui, H. Atkins & D. J. Middleton
A new genus of Gesneriaceae in China and the transfer of Briggsia species
to other genera 195
J.D. Mood, J.F. Veldkamp & L.M. Prince
A new species and a new record of Boesenbergia (Zingiberaceae)
for Thailand 207
J.D. Mood, J.F. Veldkamp, S. Dey & L.M. Prince
Nomenclatural changes in Zingiberaceae: Caulokaempferia is a superfluous
name fox Monolophus and Jirawongsea is reduced to Boesenbergia 215
L. Wijedasa, Z.Q. Shee & E. Chia
Conservation status and lectotypfication of Alangium ridleyi (Comaceae)
in Singapore 233
BOOK REVIEWS
Flora of Peninsular Malaysia. Series 11: Seed Plants, Volume 4. R. Kiew,
R.C.K. Chung, L.G. Saw & E. Soepadmo (eds). 2013. (Y.W. Low) 241
A Guide to the Native Palms of Singapore. Adrian H.B. Loo, Ang Wee Foong,
William J. Baker & Hugh TW Tan. 2014. (K.M. Wong) 243
Date of publication: 20 November 2014
Copyright ©
National Parks Board
Singapore Botanic Gardens
1 Cluny Road
Singapore 259569
Printed by Oxford Graphic Printers Pte Ltd
Gardens’ Bulletin Singapore 66(2): 121-123. 2014
121
Phlogacanthus magnus^ a new combination in the
Acanthaceae from Malaysia
Y.F. Deng
Key Laboratory of Plant Resources Conservation and Sustainable Utilization,
South China Botanical Garden, Chinese Academy of Sciences,
Guangzhou 510650, China
yfdeng@scbg.ac.cn
ABSTRACT. An examination of the type material of Gymnostachyum magnum C.B. Clarke
(Acanthaceae) from Malaysia reveals that it is a Phlogacanthus rather than a Gymnostachyum
due to having two staminodes in addition to the two fertile stamens and muticous anther-thecae.
The new combination, Phlogacanthus magnus (C.B. Clarke) Y.F.Deng is therefore proposed.
Keywords. Acanthaceae, Gymnostachyum, Malaysia, Phlogacanthus
Introduction
Gymnostachyum Nees is a genus of about 30 species of Acanthaceae distributed in the
Asian tropics (Mabberley, 2008; Flu et ah, 2011). It belongs to tribe Andrographideae
together with Andrographis Nees, Phlogacanthus Nees, Cystacanthus T.Anderson,
Diotacanthus Benth., Graphandra Imlay, Haplanthodes Kuntze and Indoneesiella
Sreem. (Lindau, 1895; Scotland, 1992; Scotland & Vollesen, 2000; Stevens, 2014;
McDade et al., 2008).
For the Malay Peninsula, Clarke (1908) recognised eleven species of
Gymnostachyum, including ten new species, and Ridley (1923) recognised 13 species.
They also suggested that Phlogacanthus and Gymnostachyum might be united
into a single genus but this is in part because they confused the circumscription of
Gymnostachyum and Phlogacanthus. Phlogacanthus, a genus of about 35 species
distributed in tropical regions of Asia, can be easily distinguished from Gymnostachyum
by having two staminodes, in addition to the two fertile stamens, and muticous anther-
thecae (Hu et al., 2011; Xia & Deng, 2013). In Gymnostachyum, the staminodes are
absent and one or both of anther-thecae are mucronate at the base. The separation
of Phlogacanthus from Gymnostachyum is also supported by molecular evidence
(McDade et al., 2008). During a visit to Singapore Botanic Gardens’ Herbarium in
2013, 1 had the opportunity to examine the type material of Gymnostachyum magnum
C.B.Clarke, one of species described by Clarke (1908). Gymnostachyum magnum has
two stamens, two staminodes and muticous anther- thecae, characters that conform
well to Phlogacanthus. Therefore, the new combination, Phlogacanthus magnus
(C.B.Clarke) Y.F.Deng, is proposed below.
122
Gard. Bull. Singapore 66(2) 2014
Phlogacanthus magnus (C.B.Clarke) Y.F.Deng, comb. nov. - Gymnostachyum
magnum C.B.Clarke, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74(3): 664 (1908). TYPE:
Malaysia, Negri Sembilan, Biikit Tampiii waterfall, May 1894, J.S. Goodenough 1893
(holotype K; isotype SING!).
Distribution and habitat. This species is endemic to Peninsular Malaysia. Its habitat
details are not well known.
Provisional lUCN Conservation Assessment. Currently, this species is only known
from three localities in Peninsular Malaysia comprising only five, mostly fairly old,
collections, but no attempt has yet been made to assess its frequency in the field. Given
the lack of knowledge of the cun*ent status of the species it should be assessed as Data
Deficient (DD) (lUCN, 2001, 2011).
Additional specimens examined: PENINSULAR MALAYSIA. Kedah: Katanh, Koh Mai
Forest Reser\^e, 4 Apr 1938, Kiah s.n. (SING). Negeri Sembilan: Sungei Ujong, Bukit Sutu,
1 Nov 1885, V.M. Alvins 1955 (SING). Selangor: Kajang, Sungei Lalauy, 9 Mar 1930, C.F.
Symington 22710 (K); ibidem, without date, C.F. Symington 22763 (SING); ibidem, 28 Mar
1931, C.F. Symington 22947 (SING).
ACKNOWLEDGEMENTS. The author would like to thank Dr Wong Khoon Meng, Mr Low
Yee Wen and Ms Serena Lee for their help during my visit to the Herbarium of Singapore
Botanic Gardens (SING). This work was supported by the National Natural Science Foundation
of China (grant nos. 31070175, 31270247, 31470302) and the Singapore Botanic Gardens’
Fellowship.
References
Clarke, C.B. (1908). Acanthaceae. In: King, G. & Gamble, J.S. Materials for a Flora of the
Malayan Peninsula. J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74: 628-698.
Hu, J.Q., Deng, Y.F, & Wood, J.R.I. (2011). Acanthaceae. In: Wu, C.Y, Raven, P. & Hong,
D.Y. (eds) Flora of China 19: 369^77. Beijing: Science Press & St. Louis: Missouri
Botanical Garden Press.
lUCN (2001). lUCN Red List Categories and Criteria, Version 3.1. Switzerland, Gland and
UK, Cambridge; lUCN.
lUCN (2011). Guidelines for using the lUCN Red List Categories and Criteria, Version 9.
http://www.iucnredlist.org/documents/RedListGuidelines.pdf (accessed on 29 Jun.
2014).
Lindau, G. (1895). Acanthaceae. In: Engler, A. (ed). Die Naturliche Pflanzenfamilien 4(3b):
274-354. Leipzig.
Mabberley, D.J. (2008). Mabberleys Plant-book: a Portable Dictionaiy of Plants, their
Classifications, and Uses. Cambridge: Cambridge University Press.
McDade, L.A., Daniel, T.F. & Kiel, C.A. (2008). Toward a comprehensive understanding of
phylogenetic relationships among lineages of Acanthaceae s.l. (Lamiales). Amer. J. Bot.
95: 1136-1152.
Phlogacanthus magnus, a new combination from Malaysia
123
Ridley, H.N. (1923). Acanthaceae. In: Ridley, H.N. (ed) The flora of the Malay Peninsula 2:
554-610. London: L. Reeve & Co.
Scotland, R.W. (1992). Pollen morphology of Andrographideae (Acanthaceae). Rev. Palaeobot.
Palynol. 72: 229-243.
Scotland, R.W. & Vollesen, K. (2000). Classification of Acanthaceae. Kew. Bull. 55: 513-589.
Stevens, P.F. (2014). [continuously updated]. Angiosperm Phylogeny Website, version 13.
http://www.mobot.org/MOBOT/research/APweb/ (accessed on 29 Jun. 2014).
Xia, C. & Deng, Y.F. (2013). Phlogacanthus yangtsekiangensis, a new combination in Chinese
Acanthaceae. Phytotaxa 104(1): 58-60.
Gardens’ Bulletin Singapore 66(2): 125-135. 2014
125
Two new species and one new subspecies of Ridleyandra
(Gesneriaceae) from Peninsular Malaysia
R. Kiew
Forest Research Institute Malaysia,
52109 Kepong, Selangor, Malaysia
ruth@frim .gov.my
ABSTRACT. Two new species, Ridleyandra bintangensis Kiew and R. nuangensis Kiew, from
Kedah and Selangor respectively, and one new subspecies, R. kiewii subsp. magnifica Kiew,
from Kelantan and Terengganu, are described from Peninsular Malaysia.
Keywords. Gesneriaceae, Kelantan, Peninsular Malaysia, Ridleyandra bintangensis, R.
nuangensis, R. kiewii subsp. magnifica, Terengganu
Introduction
One of the aims of the Flora of Peninsular Malaysia project is to explore areas that
are unknown or poorly known botanically (Kiew & Rafidah, 2007) and this has proved
fruitful in terms of the discovery of new species. Since the establishment and revision
of Ridleyandra (Weber & Burtt, 1998a, 1998b), four new species have been described
from Peninsular Malaysia, R. chuana Kiew (Kiew, 2013), R. iminii Siti-Munirah (Siti-
Munirah, 2012), R. kelantanensis Kiew (Kiew, 2009) and R. padangens is Kiew (Kiew,
2011). Here a further two species and one subspecies are described bringing the total
number of species for Peninsular Malaysia to 2 1 . Conservation assessments for these
species will be included in a forthcoming paper on conservation in the genus as a
whole.
Taxonomy
1. Ridleyandra bintangensis Kiew, sp. nov.
Among Ridleyandra species with leaves with an entire margin, inflorescences with
more than one flower, and long corollas, it most resembles Ridleyandra atropurpurea
(Ridl.) A.Weber but it differs from that species in its short petioles 1.3-2. 5 cm long
(c. 4.5 cm long in R. atropurpurea), its relatively broader oblong leaves less than
three times as long as wide (not falcate and more than three times as long as wide in
R. atropurpurea), c. 9 pairs of lateral veins (15-17 pairs in R. atropurpurea), and a
corolla that is cylindric and 10-13 mm wide at the mouth with very short lobes 2-3
mm long (trumpet-shaped and 14-18 mm wide at the mouth with lobes 5-12 mm
long in R. atropurpurea). While both species have rosy purple corollas with white
126
Gard. Bull. Singapore 66(2) 2014
lines on the lower side, the lobes of Ridleyandra bintangensis are black while those of
R. atropurpurea are concolorous with the tube. TYPE: Peninsular Malaysia, Kedah,
Sungai Selim FR (Forest Reserve), Compartment 15, trail to Gunung Bintang, 30
October 2008, Mohd. Hairul FRI 60092 (holotype KEP; isotypes SAN, SAR, SING).
(Fig. lA, 2 ).
Perennial herb. Stem woody, unbranched, erect, to 1 0 cm tall, c. 6 mm diam. Leaves in
more-or-less equal pairs, tightly clustered in a rosette at top of the stem; petioles, leaf
margin and lower surface of midrib and lateral veins densely pubescent, hairs brown,
c. 1 mm long; petioles 1.3-1.7(-2.5) cm long; lamina oblong to slightly oblanceolate,
10-13 X 3.5-5 cm, glossy, slightly bullate, glabrous above; base slightly unequal,
rounded to shortly auriculate, margin entire, wavy towards the apex, apex acute; midrib
and lateral veins flat above, prominent beneath, lateral veins c. 9 pairs. Inflorescences
axillary, (l-) 2 -flowered cyme, peduncle, bracts and calyx glabrous; peduncles deep
purple-brown, (5-)8-9.5 cm long; bracts 3, free, positioned immediately below the
flowers, reddish brown, lanceolate, 10-13 x 4 mm, apex acute; pedicel 4-13 mm
long. Sepals divided to base, reddish brown, narrowly lanceolate, 8-12 x 2-2.5 mm,
apex acute. Corolla cylindric, c. 5.5 cm to tip of lower lip; tube with a white bulbous
base, constricting to 3-A mm diam., then expanded to 10-13 mm, above the base rosy
purple with white lines on lower side, mouth glabrous; lobes 5, dull black, upper lobes
reflexed, c. 2.5 x 6 mm, lower lobes unequal, lateral lobes reflexed, c. 2 x 5.5 mm,
median lobe extended c. 2 x 5.5 mm. Stamens 4 in 2 pairs, attached c. 12 mm above
corolla base, filaments white, lower pair 13-15 mm long, upper pair 8-9 mm long;
anthers pale creamy white, c. 1.5-1 mm, without connective, staminode 1, c. 2 mm
long; nectary a naiTow white ring, c. 1-1 .5 mm high. Ovary and style 2. 7-3. 5 cm long;
stigma white, broadly triangular, c. 2.5 x 3 mm. Capsules unknown.
Distribution. Endemic in Peninsular Malaysia, known only from the type locality.
Habitat. Primary lowland dipterocarp forest at 380 m altitude.
Etymology. It is named for the type locality, Gunung Bintang, Kedah.
Notes. The flower characters that distinguish this species from Ridleyandra
atropurpurea, corolla nan*owly cylindric with black lobes vs. trumpet shaped with
rosy purple lobes, suggest different pollination syndromes but there are as yet no
observations of pollinators in Ridleyandra.
There is another species, Ridleyandra petiolata (Ridl.) A.Weber, also described
from Gunung Inas in the Bintang FR. Flowever, that species is clearly distinct from
Ridleyandra bintangensis in its petioles that are 4-8 cm long with reddish hispid hairs,
broader laminas 5-7.5 cm wide with a crenulate-serrate margin, and long reddish hispid
peduncles 15-25 cm long. According to the key in Ridley (1923) {ym^Qx Didissandra),
the corolla of Ridleyandra petiolata is violet-puiple. It was collected above 1500 m
altitude. This mirrors the Cameron Highlands where the two Ridleyandra species, R.
New Ridleyandra species and subspecies from Peninsular Malaysia
127
Fig. 1. Ridleyandra bintangensis A. Habit. R. kiewii subsp. magnifica. B. Habit. C. Corolla
opened to show stamens. D. Calyx, style and stigma. R. nuangensis E. Habit. F. infloreseence
and flower. (Photos: A, Ong P.T.; B-D, M.Y. Siti-Munirah; E-F, Yao T.L.)
New Ridleyandra species and subspecies from Peninsular Malaysia
129
morgani (Franch.) A. Weber and R. longisepala (Ridl.) A. Weber, are also separated by
altitude.
Weber & Burtt (1998b) drew attention to another specimen collected from the
Gunung Bintang range {Mohd HaniffSFN 21092) that they suggested might be a new
taxon but in its petiole 3. 5^.5 cm long, its leaves more than three times longer than
wide, its 13-15 pairs of lateral veins, and its broader corolla (14-17 mm wide) with
lobes 5-6 mm long, it is clearly similar to Ridleyandra atropurpurea (Ridl.) A. Weber.
2. Ridleyandra kiewii (Kiew) A.Weber, Beitr. Biol. Pflanzen 70: 243 (1998 [‘1997’]).
-Didissandra kiewii Kiew, Malay. Nat. J. 41 : 212 (1989). TYPE: Peninsular Malaysia,
Johor, Selai Valley, Kiew, B.H. SB 35 (holotype KEP ex UPM).
Perennial herb. Stem woody, unbranched to 20 cm long, to 7 mm diam., flowering at
c. 5 cm tall; densely hispid with femigineous hairs 4-5 mm long on the young stem,
petioles, peduncles, bracts and calyx. Leaves crowded at the apex; petioles 6-9 cm
long; lamina broadly elliptic, 10-40 x 6-18.5 cm, base cordate or rounded, sometimes
unequal, margin seiTate, crenate to undulate, apex broadly acute to acuminate, in live
state upper surface glossy dark green above, bullate and wrinkling on drying or flat,
either variegated with a broad light green band c. 2.5 cm wide along midrib or plain
green, reddish or pale green beneath, drying membranous, glabrous above, beneath
midrib shortly hispid and lateral veins minutely pubescent; lateral veins 9-18 pairs,
prominent beneath. Inflorescences axillary, cymose, either 1-3 -flowered or with up to
7 flowers; peduncles purple, 9-24.5 cm long; bracts narrowly lanceolate and 12-15 x
1-1.5 mm or foliose and 11-12 x 7-8 mm; pedicels 4-12 mm long. Calyx 10-16 mm
long, divided almost to the base, lobes 5, ligulate, 4-10 x 3-5 mm. Corolla narrowly
trumpet-shaped, outside deep purple-red or purple or tinged black, inside intensely
black, 3 .5-5 cm long, 0. 3-0.4 mm wide at base, 1-1.5 cm wide at the mouth, lobes broad
and rounded, upper two lobes more-or-less erect, 2-4 x 6-9 mm, lower three lobes 2-5
X 4-8 mm, projecting slightly beyond the upper. Stamens 4, filaments slender, white or
pale purple, lower two 19-20 mm long, upper two c. 12 mm long; anthers coimivent,
reniform, 1 .5-3 x 1-1.5 mm, cream-coloured, contrasting strongly with black corolla.
Ovary narrowly cylindric, 20-34 x 1-2 mm, glabrous, surrounded at base by a fleshy
cylindric nectary, c. 1 mm high; style narrowing to a broadly triangular pale purple
stigma c. 3 x 2.5 mm wide, stigma reaching the same level as the anthers of the shorter
stamens. Capsules glossy, dark red or purple, curved downwards at the apex, splitting
along the upper surface, 5-8 cm long, 2-6 mm wide.
Notes. It is one of the most striking Ridleyandra species with its large, purple-black
flowers and its variegated foliage. It was first discovered in Johor but recent exploration
shows that it also occurs in S Pahang, Terengganu and Kelantan. Compared with most
Ridleyandra species that are montane plants with nan'ow distributions, R. kiewii grows
in the lowlands and is the Ridleyandra species with the widest geographic distribution.
130
Gard. Bull. Singapore 66(2) 2014
The new taxon, while it is a larger plant with more serrate laminas, foliose
bracts and more flowers, resembles R. kiewii in its leaf shape, relatively long petioles,
indumentum, corolla size, shape and in its striking purple-black colour. It is treated
at subspecific rank because the two taxa are separated geographically. The species
description above is amended to take account of the new subspecies.
2a. Ridleyandra kiewii subsp. kiewii
Figure: Malayan Naturalist. 40, 2 (1986) front cover.
Lamina (10-)14(-18) x (6-)8(-9) cm, base cordate, sometimes rounded, margin
crenate to undulate, apex broadly acute, in live state upper surface bullate, sometimes
flat, variegated with a broad light green band c. 2.5 cm wide along midrib, reddish
beneath; lateral veins c. 9 pairs. Inflorescence a 1-3-flowered cyme; peduncle 9-18
cm long; bracts narrowly lanceolate, 12-1 5 x 1-1.5 mm; pedicels 4-5 mm long; ovary
c. 20 X 1 mm. Capsules 5-5.5 cm long, 2 mm wide.
Distribution. Peninsular Malaysia: Johor (Gunung Chabang Tiga, Sungai Selai) and S
Pahang (Gunung Lesong).
Habitat. Primary lowland dipterocarp forest to 400 m altitude.
2b. Ridleyandra kiewii subsp. magniflca Kiew, subsp. nov.
Compared with the typical subspecies, Ridleyandra kiewii subsp. magniflca is a
larger plant with larger leaves with more lateral veins and a more serrate margin,
the inflorescences have foliose bracts and in most cases there are more flowers per
inflorescence and the fruits are larger. TYPE: Peninsular Malaysia, Terengganu, Hulu
Terengganu, Tembat FR, Sungai Puah, 4 April 2009, Mohd. Hairul et al. FRI 60959
(holotype KEP; isotypes L, SAN, SAR, SING). (Figs IB-D, 3).
Lamina (12-)22. 5-26. 5(^0) x 11.5-18.5 cm, base rounded, sometimes cordate,
margin serrate, apex acute to acuminate, in live state upper surface flat, sometimes
bullate, most plants not variegated, sometimes variegated, especially in young
plants; lateral veins 11-1 8 pairs. Inflorescence usually a cyme with (2-)5-7 flowers,
sometimes with 1-3-flowers; peduncle (13. 5-)l 9.5-24.5 cm long, branches 1-3.5 cm
long; bracts foliose, broadly ovate, 11-12 x 7-8 mm; pedicels deep purple, 4-12 mm
long; ovary c. 34 x 2 mm. Capsules 6.5-8 cm long, 4-6 mm wide.
Distribution. Peninsular Malaysia: Kelantan (Kuala Aring), Terengganu (Telemong
FR, Tembat FR).
132
Gard. Bull. Singapore 66(2) 2014
Habitat. Primary lowland dipterocarp forest, on shaded slopes in valleys above streams,
often locally common, below 450 m altitude. In April 2009 it flowered gregariously
and in July 2006 most plants were observed with fruits suggesting that it flowers
seasonally.
Etymology. With its bunches of large, purple-black flowers, it is a magniflcent species
well worthy of cultivation and, compared with the typical subspecies, it is a larger
plant both in its leaves and fruits.
Additional specimens examined. PENINSULAR MALAYSIA: Kelantan: Kuala Aring FR,
1995, B.H. Kmx KBH 5 (KEP, E, K). Terengganu: Hulu Terengganu, Tembat FR, Sg. Puah,
1 April 2009, MoJid Hairiil et al. FRJ 60932 (KEP, SAN); ibidem, 4 April 2009, Kamarul-
Hisham et al. FBI 67096 (KEP, A, K, L, SAN, SING); Tembat FR, Sg. Tembat, 3 June 2010,
Ong et al. FRI 71064 (KEP, SING); Hulu Terengganu, Ulu Telomong, Ulu Sungai Telemong,
July 2006, R. Kiew RK 5343 (KEP, E, K, L, SAN, SAR, SING).
3. Ridleyandra nuangensis Kiew, sp. nov.
In its narrowly oblanceolate leaves and its white flowers with purple lines, it resembles
Ridleyandra wrayi (Ridl.) A. Weber, However, it is distinct from that species in its entire
leaves (dentate in Ridleyandra wrayi), its long peduncled, 2-3 -flowered inflorescences
(single flowered with peduncles to 12 cm long in R. wrayi) and the bracts that are
distant from the calyx (appressed to and enclosing the calyx in R. wrayi). TYPE:
Peninsular Malaysia, Selangor, Hulu Langat, Gunung Nuang, 21 Feb 2001, Kiew RK
5186 (holotype SING; isotypes E, KEP). (Figs lE-F, 4).
Perennial herb. Stems woody, unbranched, rarely two-branched, 30-75 cm tall, c. 5
mm diam., with a thin corky ‘bark’. Leaves in opposite pairs in a tuft at top of the
stem, below to c, 5 mm apart; petioles 1 .5-4.3 cm long; lamina oblanceolate, in life
glossy green above, whitish green beneath, glabrous, robustly succulent, 12-20 x 3.2-
6 cm, narrowed to base, margin entire, apex shortly acuminate, acumen to 5-7 mm
long; in life midrib slightly raised above, beneath scarcely prominent, lateral veins
11-13 pairs, slightly prominent beneath, midrib and veins drying dark brown beneath.
Inflorescences axillary, 2-3-flowered cymes with the flowers held high above the
leaves on peduncles 11.5-22.5 cm long; bracts and bracteoles asyimnetrically ovate,
7-10 X 3-6 mm, c. 9 mm distant 6*0111 the calyx; pedicels 0.9-2. 5 cm; flowers nodding.
Calyx pale green, 13-15 x 4-5 mm, connate for 2-3 mm, lobes lanceolate, outside
keeled, apex acute, margins shallowly toothed in the upper half, inside and out with
scattered short glandular hairs. Corolla trumpet-shaped, waxy white with a purple
line running down the centre of each lobe with the addition of fine lines in base of
throat, two white raised ridges (nectar guides) and scattered glandular hairs in throat,
minutely densely pubescent with glandular hairs outside, 4.2-6 cm to tip of lower lip;
134
Gard. Bull. Singapore 66(2) 2014
tube 3. 5-4.5 cm long, 3-7 mm diam. at base dilating to 1 1-12 mm across the mouth;
upper 2 lobes erect, 9-12 x c. 7 mm, lower 3 lobes projecthig beyond the upper, 6-10
X 6-7 mm. Stamens 4, filaments white, lower pair 1 3.5-24 mm long, upper pair 1 7-27
mm long; anthers brown, 2-2.5 x c. 2.2 mm long, joined in pairs. Nectaty annular, c.
2 mm high. Ovary and style glabrous, 17-20 mm; stigma white, broadly triangular,
4^.5 X 2-3 mm. Fruit not known.
Additional specimens examined. PENINSULAR MALAYSIA: Selangor: Hulu Langat,
Gunung Nuang, 12 Nov 2013, Yao et al. FRI 7732 9 (KEP, SAN).
Distribution. Endemic in Peninsular Malaysia, Selangor (Gunung Nuang), known
only from the type locality.
Habitat. In upper montane forest, in deep shade on slope below the summit at 1434 m
altitude.
Etymology. It is named for its type locality, Gunung Nuang, Selangor.
Notes. This species is unusual for the genus in several respects: in its tall stem (it is the
tallest species), its corky bark and its long peduncles that hold the flowers well above
the foliage.
ACKNOWLEDGEMENTS: This study is part of the research carried out for the Flora of
Peninsular Malaysia Project under ‘Safeguarding the Forest Plant Diversity of Peninsular
Malaysia’ Project No. 01-04-01-000 Khas 2 funded by the Ministry of Science, Technology
and Innovation through the National Council for Scientific Research and Development and
under ‘Dokumentasi and Inventori Flora Malaysia’ Project funded by the 10^“^ Malaysia Plan
Development Project at Forest Reseai'ch Institute Malaysia (FRIM). The project team is
grateful to the State Forestry Departments and the Department of Wildlife and National Parks,
Malaysia, for permission to make botanical collections; to the FRIM field and herbarium staff
for their assistance; to Zainal Mustafa for the botanical drawings; to Ong Poh Teck, M.Y. Siti-
Munirah and T.L. Yao for pemiission to use tlieir photographs; to Dr Gemma Bramley (K) for
providing images of Ridleyandra petiolata type specimen; to Curators of the BM, K, KLU,
SING and UKMB herbaria for permission to examine specimens in their care.
References
Kiew, R. (2009). Tliree new species of Gesneriaceae from Kelantan, Malaysia. Gard. Bull.
Singapore 61: 73-79.
Kiew, R. (2011). Two new species of Gesneriaceae from Gunung Padang, Terengganu,
Malaysia. Malayan Nat. J. 63: 661-666.
New Ridleyandra species and subspecies from Peninsular Malaysia
135
Kiew, R. (2013). Ridleyandra chuana (Gesneriaceae), a new species from Peninsular Malaysia.
PhytoKeys, 25: 15-19.
Kiew, R. & Rafidah, A.R. (2007). The Flora of Peninsular Malaysia. Conservation Malaysia.
5: 1^.
Siti-Munirah, M.Y. (2012). Ridleyandra iminii (Gesneriaceae), a new species from Peninsular
Malaysia. PhytoKeys 19: 67-70.
Ridley, H.N. (1923). Gesneriaceae. The Flora of the Malay Peninsula 2: 495-547. London: L.
Reeve & Co.
Weber, A., Burtt, B.L. (1998a [‘1997’]). Didissandra: redefinition and partition of an artificial
genus of Gesneriaceae. Beitr Biol. Pflanzen 70: 153-177.
Weber, A. & B.L. Burtt (1998b [‘1997’]). Revision of the genus Ridleyandra (Gesneriaceae).
Beitr. Biol. Pflanzen 70: 225-273.
Gardens’ Bulletin Singapore 66(2): 137-143. 2014
137
Two new white-flowered Codonoboea species
(Gesneriaceae) from Peninsular Malaysia
R. Kiew
Forest Research Institute Malaysia,
52109 Kepong, Selangor, Malaysia
ruth@frim.gov.my
ABSTRACT. Two new white-flowered Codonoboea species, C mentigiensis Kiew and C.
tembatensis Kiew (Gesneriaceae), are described from Peninsular Malaysia. Following the
lUCN Criteria and Categories, C tembatensis is Critically Endangered by clear-felling of forest
prior to the ai'ea being flooded for an extension to a hydroelectric dam, while C. mentigiensis,
that grows in lower montane forest within Pennanent Protection Forest, is of Least Concern.
Keywords. Codonoboea mentigiensis, Codonoboea tembatensis, conservation status,
Gesneriaceae, new species. Peninsular Malaysia
Introduction
Currently, 77 named species of Codonoboea are recorded for Peninsular Malaysia
(Kiew & Lim, 2011; Kiew, 2011; Kiew & Sam, 20 1 2; Lim et al., 20 1 3) and new species
continue to be discovered. The first one described below was discovered at Cameron
Highlands, Pahang, one of the best-collected localities in Peninsular Malaysia. It is
unusual among Codonoboea species in its corolla being completely white; usually
white-flowered species have contrastingly coloured (usually yellow) nectar guides.
The second species was discovered during a botanical survey of an area in Hulu
Terengganu designated for an extension to the Kenyir Hydroelectric Dam that will
be clear-felled prior to flooding. This latter species is quite unlike other Codonoboea
species, which have narrowly cylindric fruits c. 1-1 .5 mm wide, in its fruit being 3.5-5
mm thick and almost humped at the base and tapers to the tip.
Taxonomy
Codonoboea mentigiensis Kiew, sp. nov.
Codonoboea mentigiensis is similar to C. albomarginata (Hemsl.) Kiew in its few-
flowered pedunculate inflorescences and small flowers but it is different from C.
albomarginata in its larger leaves, 10-14 x 5-8 cm, that are less than twice as long
as wide (7.5-10 x 6.5 cm and 2.5-3 times longer than wide in C. albomarginata) and
in its completely white flowers 18-19 mm long (those of C. albomarginata are 12
mm long, white with a pinkish tinge due to rosy purple hairs on the outer surface, the
138
Gard. Bull. Singapore 66(2) 2014
corolla lobes are yellow and the nectar guides are orange-brown), and in its longer
fruit (3 cm not 2 cm as in C. albomarginatd). Nor does Codonoboea mentigiensis
have the purple stems, peduncles and rosy purple calyces of C albomarginata. TYPE:
Peninsular Malaysia, Pahang, Cameron Highlands, Mentigi Forest Reserve (FR), 4°
31' 48.40" N, lOR 22' 21.80" E, 18 November 2009, K. Imin FRI 68496 (holotype
KEP; isotypes K, L, SING). (Fig. 1).
Small herb. Stems 8.5-15 cm tall, 9 mm thick, semi-woody at base, pale green, with
densely appressed hairs, intemodes 2-3.5 cm long; frequently with axillary shoots.
Leaves in distant equal pairs; petioles slender, pale gi'een with downy white hairs,
3.5- 5 cm long, 3-4 mm thick, grooved above; laminas soft, pale green with a dense
velvety layer of whitish non-glandular hairs, densely softly hirsute above and beneath
especially on the midrib, margin and lower surface of veins, ovate to broadly oval,
10-14.5 X 5-8 cm, cuneate, margin entire, apex acute, lateral veins 8-11 pairs, slightly
impressed above and slightly prominent and paler beneath, intercostal veins obscure.
Inflorescences axillary, erect with a pair-flowered cyme held horizontally, peduncle
4. 5- 6. 5 cm, pale green, silky with dense glandular hairs c. 0.75 mm long; bracts
lanceolate, c. 4 x l mm. Flowers with pedicels 1.5-2 mm long; calyx mid-gi*een and
hispid with long non-glandular hairs, 5-lobed, divided almost to base, lobes naiTowly
acute, 3^(-7) x 1 - 1.2 mm long, apex slightly recurved; corolla white, 18-19 mm
long, with no markings in the throat, outside with dense long glandular translucent
hairs, tube c. 13 mm long, glabrous inside, constricted at base and c. 3 mm wide,
dilating to 5-7 mm wide at the mouth, upper lobes erect, oblong, c. 4 x 3 mm, side
lobes rounded c. 4 x 4 mm, and the lower lobe rounded c. 5 x 4 mm; the side and
lower lobes projecting 5 mm beyond the upper, with scattered minute uniseriate hairs,
stamens and stigma positioned within the mouth; stamens 2, filaments c. 3 mm long,
glabrous, straight, attached c. half-way up the corolla tube; anthers white, small, c.
1.2 mm long, joined side by side, staminodes 0; nectaiy annular, c. 0.5 mm high, rim
slightly wavy; ovary white, c. 4 mm long, ovary and style densely hispid; style white,
c. 8 mm long; stigma large, peltate, c. 1 mm across, white (not conspicuous). Fruits
horizontal, short-stalked, stalk thickened, 5-7 mm long, capsules cylindric, c. 30 mm
long, c. 2 mm wide at base, slightly upcurved, splitting only along the top, brown when
ripe.
Distribution. Peninsular Malaysia, endemic in Pahang, Cameron Highlands, Mentigi
FR.
Ecology. Lower montane forest at c. 1800-2100 m, on shaded earth banks, common
in places.
Etymology. It is named for its location, the Mentigi FR.
New Codonoboea species from Peninsular Malaysia
139
B
Fig. 1. Codonoboea mengtigiensis Kiew. A. Habit. B. Flower showing position of stamens
and pistil inside. C. Corolla opened to show position of stamens. D. Corolla cut lengthwise to
show stamens and pistil. E. Stamens showing coherence of anthers. (Drawn from FRI 68496
by Zainal Mustafa).
140
Gard. Bull. Singapore 66(2) 2014
Conset^ation Assessment. Least Concern (LC). Following the lUCN Criteria &
Categories (lUCN, 2001), although it is known from a single locality, the Mentigi FR
at Cameron Highlands, and has an AGO less 20 Ian-, it is considered to be of Least
Concern because it is lies within a Permanent Protection Forest, being a steep upland
area above 1000 m altitude and with no known threats.
Other specimens examined. PENINSULAR MALAYSIA: Pahang: Cameron Highlands,
Mentigi FR, 4 Dec 1983, Kiew RK 1263 (KEP); ibidem, 3 1 Aug 1990, RK 3119 (KEP); ibidem,
7 July 1 999, RK 4730 (KEP).
Codonoboea tembatensis Kiew, sp. nov.
Unique among Peninsular Malaysian species of Codonoboea by a combination of
its subsessile inflorescences with 1-3 flowers and short capsules (15-18 imn long)
that are unusually thick (3.5-5 mm thick) at the base. TYPE: Peninsular Malaysia,
Terengganu, Hulu Terengganu, Tembat FR, 5°12.36' N, 102° 37.10' E, 31 July 2009,
Kamarul et al. FRI 67142 (holotype KEP; isotype SAN). (Fig. 2).
Unbranched, erect herb or branching at base and forming clumps. Stems 20-45 cm
tall, 2-4 mm thick, purple with soft white glandular hairs, intemodes 1.5-3. 5 cm
long, swollen and constricted at nodes. Leaves in distant pairs; petioles slender,
lower petioles longer, 3-5.5 cm long, with soft white glandular hairs; laminas soft,
narrowly to broadly lanceolate, 9-11 x 3. 5-5. 5 cm, softly hairy (hairs not glandular),
dark green above, paler or white-green beneath, base rounded, slightly unequal and
with an additional vein on the longer side, margin shallowly crenate especially in
the upper half, apex attenuate, lateral veins (8-)9 pairs, slightly impressed above and
prominent beneath. Inflorescences axillary, a subsessile cyme with 1-3 flowers; bracts
lanceolate, 8-10 x 2-3 mm, apex rounded, midrib distinct, densely hairy. Flowers
with pale green pedicels and calyx, pedicels (2-)3^ mm long; calyx 5-lobed, 3-4(-7)
mm long, divided almost to base, lobes narrowly acute, with long hairs; corolla white,
c. 3 cm long, tube c. 20 nun long, c. 3 nun wide in the basal c. 4 nun, dilating to 13
mm wide at the mouth, upper lobes erect to reflexed, c. 7 x 9 mm, the side lobes c. 9
X 8 mm and lower lobe c. 9 x 7 mm; projecting beyond the upper, stamens and stigma
positioned within the mouth, nectar guides in the lower part of the throat consisting of
2 rows of minute, lemon-yellow, glandular hairs; stamens 2, filaments c. 10 mm long,
bowed, covered in minute, white glandular hairs, lower half of the filaments yellow,
upper half of filaments and anthers white, anthers small, c. 1 .5 mm long, joined side
by side, staminodes 0; nectaiy large, white, annular, c. 0.75 mm high; ovary white or
purple, narrowly ovoid, glabrous at base, c. 4 mm long, as long as calyx, style white
with fine, long hairs, 1 3-1 5 mm long; stigma large, peltate, c. 1 mm across, white (not
conspicuous). Fruits horizontal, short-stalked, stalk thickened, 5-7 mm long, capsules
thick at base (= humped on the upper side), 15-18 mm long, 3.5-5 mm thick at base,
tapered to apex, dull puiple becoming brown.
Fig. 2. Codonoboea tembatensis Kiew. A. Habit. B. Flo\
position of stamens and pistil. D. Calyx. E. Corolla cut ler
F. Fruit. (Drawn from FRl 67142 by Zainal Mustafa),
142
Gard. Bull. Singapore 66(2) 2014
Distribution. Peninsular Malaysia, endemic in Terengganu, Hulu Terengganu, Tembat
FR.
Ecology. In lowland to hill mixed dipterocarp forest, in valleys on shaded slopes above
streams or rivers at 200-815 m altitude. It grows as individual plants or fonns clumps
by branching at the base.
Etymology. It is named for its location, the Tembat FR.
Consen’ation Assessment. Critically Endangered, CR A3(c), Blab(i,iii,iv,v). Following
the lUCN Criteria & Categories (lUCN, 2001), this species is Critically Endangered
being at extremely high risk of extinction in the wild as its habitat lies within the area
that has been clear-felled prior to the area being flooded by the northern extension to
the Kenyir hydroelectric dam. It apparently has a restricted occurrence (less 50 km^) as
it has not been found in adjacent forests, such as the Telemong FR and the Chichir FR.
Other specimens examined. PENINSULAR MALAYSIA: Terengganu: Hulu Terengganu,
Tembat FR — Gunung Tembat, 694 m, shady valley, 10 Apr 2008 [flower], Kiew et al. FRl
57034 (KEP); Ulu Sungai Terengganu Mati, 3 Apr 2009 [fruit], Siti Munirah et al. FRI 67904
(KEP, K, L, SAN, SING); Cultivated in Forest Research Institute Malaysia’s nursery 15 Oct
2009 [flower], FRI 65159 (KEP, flowers in spirit).
Notes. The flower of Codonoboea tembatensis is unusual among Malaysian Codonoboea
in that the two nectar guides in the throat of the corolla are composed of erect, lemon-
yellow hairs (not plastids in the corolla tissues as is usual in other species). In addition,
the lower part of the filament has the same lemon-yellow colour as the nectar guides.
In most Codonoboea species the filament is uniformly white.
The fruit of Codonoboea tembatensis is relatively short (less than 2 cm long)
and is especially thick (3.5-5 mm wide) for Codonoboea species. It sometimes appears
humped at the base. Other species with fruits less than 2 cm long, like Codonoboea
floribimda (M.R.Hend.) C.L.Lim and C. reptans (Jack) C.L.Lim, have fruits 1-1.5 mm
at the base. Only Codonoboea malayana (Hook.f ) Kiew has fruits 2-3.5 mm wide but
they are longer (2.5-5 cm long). However, this latter species is not at all similar to the
new species because it has long peduncles, many-flowered inflorescences and much
longer corollas (35-40 mm long).
ACKNOWLEDGEMENTS. This study was carried out as part of the Flora of Peninsular
Malaysia Project funded by the Ministiy of Science, Technology and Innovation through the
National Council for Scientific Research and Development under Project No. 01-04-01-000
Khas 2 entitled “Safeguarding the Forest Plant Diversity of Peninsular Malaysia” and the 10'*'
Malaysia Plan Development Project entitled “Dokumentasi dan Inventor! Flora Malaysia”.
Thanks are due to the KEP herbarium staff for their support in the field, to A.R. Unnnul-Nazrah
and M.Y. Siti-Munirah for organizing the expedition to the Tembat area, to Zainal Mustafa
New Codonoboea species from Peninsular Malaysia
143
(KLU) for the preparations of the botanieal plates, and to the curators and staff of the BM, E,
K, KLU, L and SING herbaria for permission to examine specimens in their care.
References
lUCN (2001). Red List Categories and Criteria, version 3.1. Switzerland, Gland & UK,
Cambridge: lUCN Species Survival Commission, lUCN.
Kiew, R. (2011). Two new species of Gesneriaceae from Gunung Padang, Terengganu,
Malaysia. Malayan Nat. J. 63: 661-666.
Kiew, R. & Sam, Y.Y. (2012). Codonoboea personatiflora (Gesneriaceae) a new species from
Peninsular Malaysia. PhytoKeys 18: 61-66.
Kiew, R. & Lim, C.L. (2011). Names and new combinations for Peninsular Malaysian species
of Codonoboea Ridl. (Gesneriaceae). Card. Bull. Singapore 62: 253-275.
Lim, C.L., Kiew, R. & Haron, N.W. (2013). Codonoboea oreophila (Gesneriaceae), a new
species from Peninsular Malaysia. Blumea 58: 68-70.
Gardens’ Bulletin Singapore 66(2): 145-151. 2014
145
A new lectotypification of Zingiber gracile var. elatius
(Zingiberaceae)
J. Leong-Skomickova
Herbarium, Singapore Botanic Gardens,
National Parks Board, 1 Cluny Road, Singapore 259569
jana_skoraickova@nparks.gov.sg
ABSTRACT. The previous lectotypification of Zingiber gracile Jack, var. elatius Ridl. is
deemed ineffective and a new lectotypification is here proposed. The original varietal epithet
'elatior\ later also used at specific rank, is corrected here to 'elatius', to agree grammatically
with the generic name.
Keywords. H.N. Ridley, lectotypification, Penang Hill, Zingiber elatior, Z. elatius, Z. gracile
Introduction
During my recent work on Zingiber singapurense Skomick., a new species from
V
Singapore (Leong-Skomickova et al., 20 1 4), all of the relevant literature on the Zingiber
gracile complex was studied, including the protologues and existing revisions, along
with an examination of the type specimens.
Zingiber elatius (Ridl.) Theilade was originally published and repeated in
subsequent works with an incorrect grammatical Latin termination as Zingiber gracile
var. elatior! Zingiber elatior^ but is coiTected here to 'elatius' following Arts 23.5, 24.2
and 32.2 of the ICN (McNeill et al., 2012). When discussing earlier work 1 use the
spelling ‘elatius’, even when the authors of those earlier works used ‘elatior’, unless
I am quoting directly. It is currently recognised at specific rank, a conclusion with
which I ftilly concur, but I discovered that there is nomenclatural confusion around
this name. My initial failure to locate the specimen at SING that had been designated
by Theilade (1998) as the lectotype of 'Zingiber elatior Ridl.’ (= Zingiber gracile
Jack var. elatius Ridl.) led not only to the realisation that Ridley never recognised this
name at specific rank, but also to a number of bibliographic mistakes in Theilade ’s
entry under this taxon and lectotype designation. The lectotype she designated
is not an actual specimen but rather a mixture of data (locality, date and collection
number) from two unrelated specimens representing two different taxa. Tlie history
of the basionym. Zingiber gracile var. elatius, the original material connected to it,
and previous comments on this issue are chronologically reviewed in this paper. The
reasons why previous lectotypification have to be deemed ineffective are explained,
and the new lectotypification is proposed.
146
Card. Bull. Singapore 66(2) 2014
Ridley’s brief entry (1899) on Zingiber gracile var. elatius reads [verbatim,
including the spelling and formatting]:
Var. elatior
A very much taller slenderer plant with stems about five
feet tall, leaves narrow linear lanceolate acuminate 10 inches by
1, peduncle 18 inches and spike 7 or 8.
Hills at 2000 feet or upwards. Penang Hill. Perak, Max-
well’s Hill.
King No. 7954. Possibly a distinct species.
Ridley cited material from tliree locations, (1) Penang Hill, (2) Perak, Maxwell’s Hill,
and (3) King No. 7954 [collected from Perak], albeit with a remark that it is possibly
a distinct species. Although Furtado noted (7 September 1931, handwritten note on
one of the two sheets of King 7954 in SING with an almost identical note also on
the duplicate at K) “T take that Ridley used the words ‘possibly a distinct species’ to
qualify his var elatior and all the specimens quoted under it, not this specimen only”,
it is possible (as suggested by the fomiatting) that Ridley’s remark belongs exclusively
to the last element (3) King 7954 which, unlike the elements (1) and (2), occurs in the
lowlands (300-500 ft as per details on the specimens). In fact, when comparing Ridley’s
eligible original material from these three locations it is possible that three taxa are
involved. Element (1) [single sheet at K, K000255246] from Penang Hill is certainly
a different taxon from elements (2) [single sheet at SING, SINGO 176950] and (3) [2
sheets at SING, SING0176448, SINGOl 76449; 1 sheet at K, K000255245]. Elements
(2) and (3), although somewhat more similar to each other than to element ( 1 ), are also
likely to be two different taxa based on inflorescence shape and proportions. Ridley’s
brief description of his Zingiber gracile var. elatius is a fusion of leaf measurements
from element ( 1 ) and inflorescence measurements from element (2), while element (3)
does not match the description in both leaves and inflorescence dimensions.
Holttum (1950) upheld this taxon at varietal level but did not select or indicate a
preference for any of the original material as a type. Holttum ( 1 950) accepted Furtado ’s
interpretation of Ridley’s entry, i.e. that Zingiber gracile var. elatius is possibly a
distinct species, a view that was later shared by Theilade (1998) who went further, and
incorrectly, interpreted this statement as Ridley’s definitive acceptance of this taxon at
specific level.
Theilade (1998), while revising Zingiber for Peninsular Malaysia, recognised
all four varieties of Zingiber gracile listed by Holttum {Zingiber gracile var. gracile,
Z. gracile var. aurantiacum Holttum, Z. gracile var. elatius Ridl. and Z. gracile var.
petiolatum Holttum) at specific rank. For two of these, Zingiber aurantiacum (Holttum)
Theilade and Zingiber petiolatum (Holttum) Theilade, she was quite explicit about
raising the taxa to the rank of species and also designated lectotypes.
Theilade (1998) indicated Ridley as the author for Zingiber elatius even though
Ridley (1899) clearly described this taxon at the varietal level and never published this
A new lectotypification of Zingiber gracile var. elatius
147
name at specific rank. She also selected a lectotype for this name, making it clear she
did not consider herself to be the one effecting the change of rank.
Theilade’s nomenclatiiral section reads [verbatim; discrepancies underlined]:
Zingiber elatior Ridl. 11899k J. Straits Brch. R. Asiat. Soc. 32: 130; (1924)
FI. Mai. Pen. 4: 260. — Z. gracile var. elatior Holttum (1950), Gdns’ Bull.
Singapore 13: 64.
There are two problems with this entry. Firstly, Ridley never published Zingiber
elatius at specific rank in either of the two works cited by Theilade (i.e., Ridley,
1899, 1924). Secondly, the basionym should be Zingiber gracile Jack. var. elatius
Ridl. but, for unknown reasons, Theilade assigns this name in her citation to Flolttum
(1950). Holttum, however, never claimed this name as his own but clearly assigned
it to Ridley (Holttum, 1950: 64). Theilade’s mistake in assigning the authorship of
Zingiber elatius to Ridley was highlighted by Turner (2000) who suggested that the
con'ect authorship should be Zingiber elatius (Ridl.) Theilade. Turner (2000) noted
that ‘notwithstanding several en'ors in bibliographic citation, the use of elatior is
admissible as the publication of a new combination’. Turner’s approach is in line with
Art 41.6 of the ICN (McNeill et ah, 2012) which states that errors in the citation of a
basionym, including an incorrect author citation, do not preclude valid publication of
a name at a new ranlc.
The designation of the lectotype of Zingiber elatior is equally conflising
(Theilade, 1998) and reads [verbatim; discrepancies underlined]:
Type: Peninsular Malaysia, Penang. Apr. 1896, Ridley 9340 (SING lectotype!,
K isotype!) Lectotype selected here.
The major problem with this statement is that no herbarium specimen exactly matching
the above citation exists in SING or K.
The SING herbarium does not have, and has no record of ever having had, a
specimen of Ridley 9340, contrary to Theilade’s and Turner’s statements of it being in
SING. This has been confinued by a manual search of the entire Zingiber collection as
well as the entire Zingiberaceae type collection. Moreover, the entire general collection
of the Zingiberaceae at SING has been barcoded and databased and Ridley 9340 is not
on record. There was also no sign of a Ridley 9340 specimen in the now discontinued
but still extant card index at SING which contains all of Ridley’s collections in SING.
From the 28 entries in the card index covering Ridley’s collection numbers from 9226
to 9378 (including the closest numbers 9338 and 9342), it is obvious that Ridley
collected in Penang in June 1898. This implies that Ridley 9340 would also have been
collected in June 1 898. This conflicts with Theilade’s information, which dated Ridley
9340 to April 1896. The sheet with the closest information to Theilade’s citation at
SING is a specimen with barcode number SO 176947 (Fig. 1), collected by Ridley
in April 1896 in Penang, but with no collection number and no exact locality given.
cm copyright reserved
148
Gard. Bull. Singapore 66(2) 2014
MALAY PENINSULA. ** " /■
>■ ‘''-VAXG. ^ a. -
\itfive Xame ' ^ **
Dotiiuicnl. <w ,.■
/.oi (/ /i/jf iiiPrtV
EtevaUo,,.:,
Fig. 1. Specimen of Zingiber gracile Jack. (SING; SOI 76947) collected by Ridley in Penang,
April 1896. Reproduced with permission of Singapore Botanic Gardens.
A new lectotypification of Zingiber gracile var. elatius
149
There is no mention of Penang Hill - a locality which Ridley cites in his original entry.
Penang Hill (823 m), being the highest point of Penang Island, is a more specific
locality with a distinct flora than a generic Penang which often refers to lowland
locations elsewhere on the island. Moreover this sheet is not a specimen of Zingiber
gracile var. elatius, as outlined by Ridley, but rather Z. gracile var. gracile for which
the type locality is also in Penang (note the ovate to elliptic leaves, long papery ligules
and short peduncle). Holttum (1950), in fact, correctly cited this particular specimen
under the nominal variety of Zingiber gracile in his account. What possibly confused
Theilade is an addition to Ridley's original label in an unknown hand of ‘var. elatius'
and another remark in Furtado’s hand that this sheet is the syntype of the variety (see
Fig 1). Tt should also be noted, however, that there is no annotation by Theilade that
would clearly indicate that this is indeed the specimen she intended to designate as
the lectotype. Nevertheless, this specimen should not be considered to be part of the
original material of Zingiber gracile var. elatius as it is not a good fit to the protologue
in both locality data (as explained above) and morphology (it does not match the
original description in either inflorescence or in leaf dimensions with peduncles c, 4
inches long, spike 4 '4 inch long and largest lamina c. 6 x c.\% inches compared to 18
inches long inflorescence, 7-8 inches long spike and lamina 10x1 inches as stated in
the protologue).
The Kew herbarium has a single sheet of Ridley 9340 [K000255246] in its
holdings (Fig. 2). As expected, based on the search of the SING card index, the collection
date is indeed June 1 898 and the locality is recorded as Penang Hill, Richmond Pool,
2500 ft. This specimen is one of the two sheets on which Ridley based his description.
Moreover, Ridley cited a collection from Penang Hill and this specimen is the only
collection of Zingiber from Penang Hill which fits the original description at least
partly (as explained in the beginning of the introduction) and predates the publication
of the name.
In view of the information given above, Theilade ’s lectotypification has to
be considered ineffective because a particular specimen was not clearly indicated as
a lectotype and is, therefore, contrary to the ICN Art. 7.10 (McNeill et ah, 2012).
The Ridley specimen at SING (barcode SINGOl 76947) is not annotated as Ridley
9340 and does not fit the original description. On the other hand, it seems obvious
that Theilade’s “isotype” is the Ridley 9340 at K (barcode K000255246), even though
this belongs to a different gathering. In the absence of independent evidence, such as
Theilade’s annotations on either of the two specimens, it is impossible to make sense
of Theilade’s type designation and to make any correction under Art. 9.9 (McNeill
et ah, 2012). Ridley 9340 (K) [K000255246] is, therefore, selected below as the new
lectotype of the name Zingiber gracile Jack. var. elatius Ridh
While Turner (2000) spotted discrepancies in the date and locality in Theilade’s
designation of Ridley 9340 as a lectotyj^e, and corrected it in his paper to June 1898,
Penang Hill, he unfortunately did not realise that this collection is not extant at SING.
Turner (2000) also considered the specimen of Zingber gracile var. gracile discussed
above, Ridley s.n. from Penang dated April 1896 (barcode SOI 76947), to be a syntype
of Z gracile var. elatius. Tliis specimen is not, however, part of the original material.
150
Gard. Bull. Singapore 66(2) 2014
HERB. HORT, KEW.
A
rULitU I'EX’A.VG.
Nflih'e XfiBie
[AKaiiiy.^
Ekr^iatlon . , Dale
HERB. HORT, EOT.
,H . ^3>V0
SyMTVI’E
ROYAL BOTANIC QARDENS KEW
llllllliiilili
I K005255246
Fig. 2. Specimen of Zingiber elatius (Ridl.) Theilade (K; K000255246) collected by Ridley in
Penang Hill, in 1 898 and designated here as the lectotype. Reproduced with permission of the
Director and Board of Trustees, Royal Botanic Gardens, Kew.
KODAK Color Control Patches
A new lectotypification of Zingiber gracile var. elatius
151
Notes on the identity of Zingiber elatius from the type locality on Penang Hill, along
with a colour plate, can be found in Leong-Skomickova et al. (2014). Confinnation
of the identities of the remaining two elements from Ridley’s protologue are pending
re-collection of flowering material from their respective localities.
New lectotypification
Zingiber elatius (Ridl.) Theilade, Gard. Bull. Singapore 48: 227 (1998 [‘1996’]). -
Zingiber gracile Jack var. elatius Ridl., J. Straits Branch Roy. Asiat. Soc. 32: 130
(1899); Ridley, FL Malay Penins. 4: 260 (1924); Holltum, Gard. Bull. Singapore 13:
64 (1950). TYPE: Peninsular Malaysia, Penang, Penang Hill, Richmond Pool, 2500 ft,
June 1898, Ridley 9340 (lectotype K! [barcode K000255246], designated here).
ACKNOWLEDGEMENTS. I thank Siti Nur Bazilah Ibrahim (SING) for her help in our
extensive search for Ridley 9340 at SING as well as for scanning the specimen depicted in
Fig. 1; Singapore Botanic Gardens for pennission to reproduce the image of the specimen in
Fig. 1; and the Director and Board of Trustees, Royal Botanic Gardens, Kew for pennission
to reproduce the image of the specimen in Fig. 2. I also thank Prof. Karol Marhold and Mr
Otakar Sida for constructive comments on the earlier version of this manuscript. Prof John
McNeill and an anonymous reviewer are thanked for critical feedback suggesting useful
improvements to this paper including bringing to our attention the error on Latin termination of
the specific/varietal epithet. Dr David Middleton is acknowledged for language improvements
and navigating this manuscript through reviews.
References
Holttum, R.E. (1950). The Zingiberaceae of the Malay Peninsula. Gard. Bull Singapore 13:
1-249.
Leong-Skornickova, J., Thame, A. & Chew, P.T. (2014). Notes on Singapore native Zingiberales
1: A new species of Zingiber and notes on the identities of two further Zingiber taxa.
Gard. Bull. Singapore 66(2): 153—167.
McNeill, J., Barrie, F.R., Buck, W.R., Demoulin, V., Greuter, W., Hawksworth, D.L.,
Herendeen, P.S., Knapp, S., Marhold, K., Prado, J., Pmd‘homme van Reine, W.F., Smith,
G.F., Wiersema, J.H. & Turland, N.J. (2012). International Code of Nomenclature for
algae, fungi and plants (Melbourne Code). Regnum Vegetabile 154. 205 pp. Konigstein:
Koeltz Scientific Books.
Ridley, H.N. (1899). The Scitamineae of the Malay Peninsula. J. Straits Branch Roy. Asiat.
Soc. 32: 85-184.
Ridley, H.N. (1924). Zingiberaceae. In: Ridley, H.N. (ed) The Flora of the Malay Peninsula 4:
233-285. London: L. Reeve & Co.
Theilade, I. (1998 [‘1996’]). Revision of the genus Zingiber in Peninsular Malaysia. Gard.
Bull. Singapore 48: 207-236.
Turner, I. (2000). The plant taxa of H.N. Ridley, 3. The Zingiberales. Asian J. Prop. Biol. 4(1):
1-17.
Gardens’ Bulletin Singapore 66(2): 153-167. 2014
153
Notes on Singapore native Zingiberales I:
A new species of Zingiber and notes on the
identities of two further Zingiber taxa
J. Leong-Skomickoval, A. Thame^ & P.T. Chew^
^Herbarium, Singapore Botanic Gardens,
National Parks Board, 1 Cluny Road, Singapore 259569
j ana_skomicko va@nparks. gov. sg
^Conservation Division, Central Nature Reserve,
National Parks Board, 1 Cluny Road, Singapore 259569
ABSTRACT. A new species of Zingiber singapurense Skomick., is described and illustrated
here. It is compared to other species from the Zingiber gracile {Zingiber sect. Zingiber)
alliance. A key to this group of species in Peninsular Malaysia and Singapore and a key to all
Singapore native and naturalised Zingiber species are provided. The national and international
conservation status of the new species and a conservation strategy for it are discussed. Notes
on the varietal identity of Zingiber ptiberulum and the presence of Z. ottensii, previously
misidentified as Z. zerumbet, in Singapore are given.
Keywords. Central Catchment Nature Reserve, conservation, lUCN, Zingiber gracile, Z.
griffithii, Z. ottensii, Z. puberulum, Z. singapurense, Z. zerumbet, Zingiberaceae
Introduction
A conservation project on Singapore’s native Zingiberales was initiated by the Research
& Conservation Branch of the Singapore Botanic Gardens, National Parks Board,
in January 2011. Although the initial 15 months of funding has expired, the project
continues in various forms with support from a wide network of collaborators within
and outside NParks. Initially, the project mainly involved intensive surveys of forested
areas, marking of all existing populations/mature adult plants of native Zingiberales,
regular monitoring of flowering and seed-set, and ex situ and in vitro propagation
of native material from Singapore stock for re-introduction puiposes. These efforts
continue but, as we progress with the project, we are also realising that there is a need
to clarify various taxonomic issues (correct identitifications, nomenclature). We are
also conducting in-depth studies of reproductive biology and are updating the lUCN
conservation status for all native Zingiberales in Singapore. This paper is the first in
the series entitled ‘Notes on Singapore Native Zingiberales’. This series will include
rediscoveries of taxa presumed to be extinct in Singapore, provide taxonomic and
nomenclature updates, and raise awareness of, and encourage an active approach
towards, the conservation of native Zingiberales and other plants in Singapore.
154
Gard. Bull. Singapore 66(2) 2014
Information on Singapore’s geography and vegetation types, and the affinities
of its flora to those of Peninsular Malaysia and Sumatra, have recently been discussed
in detail (e.g. Chong et al., 2011; Ng et al, 2011; Yee et ah, 2011; Low et al., 2014)
and are, therefore, not repeated here. Recently, as interest has increased, numerous
surveys have reported new records for Singapore as well as rediscoveries of presumed
locally extinct species (e.g. Rodda et al., 2012; Yeoh et al., 2013; Low et al., 2014).
As Singapore’s flora is comparatively well known, and forested areas with primary
and mature secondary forests fairly small (accounting for about 3% - Yee et al., 2011),
reports of a new plant taxon from Singapore are infrequent. A Cry’ptocoryne Fisch. ex
Wydler hybrid from Bukit Timah Nature Reserve was reported in 2001 (Bastmeijer
& Kiew, 2001). Nine new species of Thottea Rottb. were also described by Yao
(2013) from Peninsular Malaysia, of which T. praetermissa T.L.Yao is also found in
Singapore. The most recent example is of two Utania G.Don species (Gentianaceae)
occurring in Singapore and the southern part of Peninsular Malaysia (Sugumaran &
Wong, 2014), based on types from Singapore. The presence of the Thottea and the two
Utania species were, however, already recorded in Singapore from the 1890s with
multiple collections for all thi'ee taxa, albeit misidentified. In tins paper, we present a
new ginger species for which no earlier collections are known.
In 2012, two sterile living specimens of a small Zingiber Mill, species were
collected during a regular survey in Singapore’s Central Catchment Nature Reseiwe by
William Ng of the National Parks Board, Singapore. Unfortunately, neither of the plants
survived and the original colleetion locality, said to contain just a few individuals,
could not be re-located. In October 2013 and April 2014, the same species was found
in three different locations within patches of primary and mature secondary forest in
the Central Catchment Nature Reserve. In May 2014, flowering plants were found at
all three locations. These collections could not be matched to any Zingiber species
from Singapore, Peninsular Malaysia, Borneo or Sumatra after careful study of type
material, additional herbarium specimens and the protologues of all related species of
Zingiber sect. Zingiber from the Malesian region by the first author. It was, therefore,
concluded that these new collections belong to a taxon as yet unknown to science which
we here name Zingiber singapurense. The description and measurements are made
from living material from all tliree populations. As previous works by Holttum (1950)
and Theilade (1998) were mainly based on herbarium specimens, the dimensions of
certain parts from dried material are also given to allow direct comparison to their
descriptions and to type material of related taxa.
Taxonomy
Zingiber singapurense Skomick., sp. nov.
Similar to Zingiber aurantiacum (Holttum) Theilade, but differing in the smaller and
more slender habit, inflorescences with distinctly convex fertile bracts (versus tightly
appressed bracts), reduced bracteoles (up to 6 mm long versus c. 25 mm long) and
Singapore native Zingiberales I.
155
lateral staminodes partly free from labellum (versus staminodes almost fidly connate
with labellum). TYPE: Singapore, Central Catchment Nature Reserve, MacRitchie
sector, 7 May 2014, J. Leong-Skornickovd & Aung Thame SNG-178 (holotype SING
(including spirit material); isotypes E, K, KEP). (Fig. 1, 2)
Perennial evergreen rhizomatous herb to 0.8 m tall. Rhizome 7-10 mm in diam.,
branched, internally light yellow, mildly aromatic. Leafy shoots of mature flowering
individuals 0.7-1. 5 m long, strongly arching, with 18-30 leaves; basal Va - Vs leafless,
with 4-5 green glabrous sheathing bracts; leaf sheaths green, glabrous; ligule 1-2 mm
long, almost inconspicuous, bilobed, green, tomentose; petiole reduced to pulvinus,
c. 2-3 mm long, light green, tomentose; lamina to 1 5-22 x 2-A cm (largest leaves
of mature flowering individuals measured; dimensions when dried 14-20 x 1.8-2. 8
cm), narrowly ovate, gradually tapering to a naiTowly attenuate-nan'owly caudate
apex, base obtuse to rounded, above dark glossy green and glabrous, lighter green
and glabrous beneath (except occasional space hairs near the midrib and the basal part
near pulvinus). Inflorescence radical, to 23 cm long, erect; peduncle 5-10(-12) cm
(comprising half of the length of the entire inflorescence); sheathing bracts 4-6, 0.5-3
cm long (basalmost shortest at c. 5 mm, gradually longer towards the spike), tubular
at base (2-10 mm), cream-white to light green (those below the leaf litter layer cream-
coloured, those above ± light green), with orange-reddish tinge in upper bracts, softly
pubescent (hairs appressed); spike 7-1 1x1 .5(-2) cm broad (dimensions when dried 7-1 1
X 1-1.3 cm), narrowly fusiform, consisting of 8-17 bracts (lowermost and uppermost
2 bracts usually sterile); fertile bracts slightly obovate, lower bracts up to 3.6 x 2.3
cm (gradually smaller towards the apex), with broadly acute to obtuse apex, externally
cream- white to light yellow at base, richly tinged orange-red in upper exposed parts,
softly pubescent (hairs appressed), internally light yellow, glossy, glabrous, margin c.
0.5 mm, hyaline, translucent white, enclosing single flower; bracteole much reduced,
to 2-7 mm long, 2-5 mm wide at base, triangular, translucent white, externally with
sparse soft white appressed hairs, internally glabrous. Flower 6-7 cm long; calyx 13-
20 mm long, cylindric in basal third, inflated in apical part, with three inconspicuous
teeth, incised unilaterally 6-10 mm, translucent white, sparsely pubescent towards
the ovary, glabrous in apical half; floral tube 35-42 mm long, cream-white at base,
yellowish towards apex; dorsal corolla lobe ovate-triangular, 26-28 x 9-1 1 mm,
translucent pale yellow, glabrous, apex narrowly acuminate, with indexed margins,
rarely minutely cuculate; lateral corolla lobes narrowly ovate-triangular, 23-25 x
5-7 mm, translucent pale yellow, glabrous, apex narrowly acuminate, with inflexed
margins; labellum 21-23 mm long, 9-1 1 mm wide (18-20 imn wide inclusive of the
lateral staminodes which are connate to the labellum and form a trilobed structure),
pale yellow, glabrous; lateral staminodes 13-14 x 4-5 mm, connate to labellum in
basal Vs, pale yellow, glabrous. Stamen 18-20 mm long (without straightening out
the anther c\:qsX)\ filament 1-1.5 mm long; anther c. 11 x 4 mm (excluding the crest),
connective tissue cream-white, anther crest 11-13 mm long (straightened), exceeding
stigma by 1-3 mm, cream- white to pale yellow at base, slightly darker towards apex;
156
Gard. Bull. Singapore 66(2) 2014
Fig. 1. Zingiber singapurense Skomick. A. Habit. B. Inflorescence. C. Rooting plantlet arising
from the apical part of the leafy shoot. D. Detail of ligule. From type SNG-178. (Photos: Jana
Leong- Skomicko va)
Singapore native Zingiberales I.
157
— ^ — —
Fig. 2. Zingiber singapurense Skornick. A. Flower (side view). B. Flower (front view). C.
Flower dissection (from left): Floral tube with ovary, calyx and anther attached, dorsal corolla
lobe, two lateral corolla lobes, labellum with connated lateral staminodes, bracteole, fertile
bract. D. Detail of ovary with epigynous glands, anther (side view) and anther (front view).
From type SNG-178. (Photos: Jana Leong-Skornickova)
anther thecae dehiscing along entire length. Epigynous glands two, c. 3.5 mm long,
c. 0.75 mm in diam., cream-white. Ovary 3^ x 2-3 mm, cream-white, pubescent,
hairs light rusty brown; style white, stigma with round, forward-facing ciliate ostiole.
Fruits not seen.
158
Gard. Bull. Singapore 66(2) 2014
Ecology and phenologyK Zingiber singapurense occurs in primary and mature secondary
forest, preferring moist and shady conditions. Flowering in May-June.
Distribution. So far endemic to Singapore.
Etymology. The specific epithet of this small and vulnerable yet resilient species is
derived from Singapore’s name in Malay, ‘Singapura’.
Additional specimens examined. SINGAPORE: Central Catclnnent Nature Reserve, MacRitchie
sector, 21 May 2014, J. Leong-Skornickovd &Aung Thame GRC-l 79 (E, SING); 29 May 2014,
MacRitchie sector, J. Leong-Skornickovd & Aung Thame GRC-l 80 (E, SING);
Provisional lUCN conservation assessment & consei^ation strategy^. So far only three
populations in the Central Catchment Nature Reserve have been discovered with each
population only having 10-25 mature individuals and less than 50 mature individuals in
total. The Extent of Occurrence (EOO) is 0.06 km^. Eligh visitor numbers and proposed
development plans could adversely affect the quality of the habitat. Following the
lUCN criteria (lUCN, 2012) and the criteria for national conservation assessments as
outlined by Davison (2008), Zingiber singapurense should be considered nationally
and globally Critically Endangered (Blab(iii,iv,v); D).
Although the leafy shoots of most Zingiberaceae are pseudostems composed
of leaf sheaths, true stems with buds, capable of producing bulbils or seedlings,
have been reported in some Globba and Zingiber species. The stems of Zingiber
singapurense are arching, a feature more prominent in older stems. The tips eventually
touch the ground and new plantlets grow from the buds (Fig. 1C), This capability to
reproduce vegetatively makes Zingiber singapurense less dependent on the presence
of pollinators and is certainly a reason behind its resilience, Flowever, it does appear
that the populations also reproduce from seeds (assumed from the presence of small
independent plantlets not physically connected to adult individuals). Nothing is
cun'ently known of the genetic diversity in the three existing populations although such
work is planned in the near future. Work on DN A barcoding of all native Zingiberales
is also in progress (Khew et al., unpublished).
The existing populations will continue to be closely monitored for fruit-set
and the numbers of seedlmgs and vegetatively produced offsprings. By selective
harvesting of young plantlets from overcrowded parts of the populations the chances
of survival of the remaining plantlets in the field improves. The harvested plantlets will
be replanted in Singapore Botanic Gardens as stock for ex situ conservation. Currently,
about 15 plants have been successfully established in the ex situ native gingers
geraiplasm collection at the Singapore Botanic Gardens. These are individuals from
all three populations to retain as much of the original genetic variation as possible.
The establishment of cuttings from stem buds and of in vitro propagation to bulk up
the number of plants is also in progress. These should produce enough stock for re-
introductions into suitable habitats as well as for its introduction into the horticulture
trade to lower the risk of illegal harvesting from the wild populations.
Singapore native Zingiberales I.
159
Notes. Zingiber is the largest genus in the sub-family Zingiberoideae with more than
200 names coiTesponding to approximately 100-150 species (Wu & Larsen, 2000;
Kishor & Leong-Skomickova, 2013). The genus is well characterised by lateral
staminodes connate to the labellum, an elongated hom-like anther crest wrapped
around the stigma, and the presence of a pulvinus. Zingiber has been traditionally
divided into four sections based on the position of the inflorescence {Zingiber sect.
Zingiber, sect. Dymczewiczia (Horan.) Benth., sect. Pleuranthesis Benth. and sect.
Cry’ptanthium Horan.). While a recent molecular study based on a single marker and
limited material hints i\\3t Zingiber sect. Dymczewiczia and Zingiber sect. Pleuranthesis
are not well segregated from Zingiber sect. Zingiber (Theerakulpisut et al., 2012), the
old classification remains in use until a new classification is formally proposed.
In its general habit and the position of the inflorescence, Zingiber singapurense
falls clearly into Zingiber sect. Zingiber. In its narrowly fusifonn orange to red
inflorescences and pale yellow flowers, it can be further narrowed to the group of
species in the Zingiber gracile alliance (for a key to Zingiber in Peninsular Malaysia see
Theilade, 1998), The complexity of this group in Peninsular Malaysia was previously
noted by Holttum (1950), who recognised four varieties (Zingiber gracile van gracile,
Z. gracile van aurantiacum Holttum, Z. gracile van elatius Ridl. and Z. gracile van
petiolatum Holttum). Theilade (1998), who revised the genus Zingiber for Peninsular
Malaysia, upgraded all varieties to the specific level (Z. gracile Jack, Z. aurantiacum
(Holttum) Theilade, Z. elatius (Ridl.) Theilade and Z. petiolatum (Holttum) Theilade).
Theilade also described an additional species fi’om this group. Zingiber suphureum
Burkill ex Theilade (Cowley & Theilade, 1995). Lim (2003) added another two species
with nan*owly fusiform inflorescences to this group, Zingiber raja C.K.Lim & Kharuk.
and Z. kelantanense C.K.Lim.
Of the seven species in the Zingiber gracile alliance, four have elliptic to ovate
laminae with an approximate length:width ratio of < 4.5. The large Zingiber raja
(type from Perak, Belum F.R.), with pseudostems to 2.5 m tall and elliptic laminae
(lengthiwidth ratio 3.6-4.23; based on the protologue), can be excluded on account of
its very long inflorescences (peduncles to 62 cm, spikes to 36 cm), markedly distinct
dark maroon labellum, and staminodes with yellow blotches. Zingiber kelantanense
(type from Kelantan, Stong F.R.) is also a large species to 2.5 m in height, with
elliptic, plicate and prominently petiolate laminae (length:width ratio 3^.1; based
on the protologue). The following two species are smaller in habit to 1 m in height.
Zingiber gracile (type from Penang), stands out in this gi'oup due to its long, papery,
thin ligules (up to 1,8 cm in the type herbarium material) and elliptic lamina (to 16 x
4.2 cm, length: width ratio 3.8; based on the lectotype) and well-developed bracteoles
(c. 20-25 mm long). Zingiber sulphureum (type from Taman Negara) differs in its
elliptic to ovate, abaxially hairy lamina (to 15^5 cm, length: width ratio 3; based on
an isotype), sulphur yellow bracts at anthesis, prominently bilobed papery ligules to 6
mm long and completely missuig bracteoles.
Zingiber singapurense has dark green, narrowly ovate to naiTOwly elliptic
laminae (14-20 x 1 .8-2.8 cm, length:width ratio 7. 1-7.8; based on herbarium material
of the holotype and isotypes) and short ligules, therefore in general habit similar to the
160
Gard. Bull. Singapore 66(2) 2014
remaining three species. Delimitation of Zingiber singapurense within this group is,
however, straightforward as it is the only species which has much reduced triangular
bracteoles (to 2-7 mm long) and distinctly convex bracts, giving the inflorescence a
bullate appearance (Fig. IB). It is also the smallest and most delicate species in the
group, with leafy shoots to 1.5 m long, but distinctly arching and not exceeding 0.8
m in height. The bracteoles are well developed in Zingiber aurantiacum (c. 25 mm),
Z elatius (25-30 mm) and Z. petiolatum (c. 30 nmi). Zingiber petiolatum (type from
Kedah) is a much larger species, up to 3 m tall, with somewhat petiolate leaves up
to 34/40 X 5.5 cm, and inflorescences up to 86/110 cm long, inclusive of the spike to
34/44 cm long (measured from the lectotype/isolectotype; lamina length: width ratio
6.2/7.2). Zingiber elatius (type from Penang Hill) is clearly distinct in the inflorescence
composed of a long peduncle (to 36 cm; in type material), an ovoid spike composed
of bronze to red-purple bracts (c. 11 x 2.6 cm; in isolectotype), and flowers with a
pale yellow label him ornamented with fine dark purple lines and tinged in the centre.
The lamina length: width ratio is c. 10 (Fig. 3 A, B). Zingiber aurantiacum (type from
Pahang, Fraser’s Hill), is a more robust, montane species up to 2 m tall, with larger
leaves (to 26 x 4.1 cm, length:width ratio c. 6.2; based on the lectotype), reddish leaf-
sheaths in young shoots, larger inflorescences (to 43 cm in type material) with tightly
appressed bright orange fertile bracts at anthesis, and lateral staminodes almost fiilly
connate with the labellum (Fig. 3C, D).
Key to species from the Zingiber gracile alliance
in Peninsular Malaysia and Singapore
la. Lamina elliptic to ovate (length:width ratio < 4.5) 2
lb. Lamina narrowly ovate to narrowly elliptic (length:width ratio >6) 5
2a. Ligules short (c. 2 mm), labellum and staminodes dark maroon with yellow
blotches Z raja
2b. Ligules 4-22 mm, flowers light yellow 3
3a. Plant to 2.5 m tall, lamina petiolate, prominently plicate Z kelantanense
3b. Plant to 1 m tall, lamina ± sessile, smooth or weakly bullate 4
4a. Ligules to 18 mm, fertile bracts orange to pink at anthesis, bracteoles well
developed Z gracile
4b. Ligules 4-6 mm, fertile bracts sulphur yellow at anthesis, bracteoles absent
Z sulphureum
5a. Bracts prominently convex, bracteoles reduced (to 7 mm long)
Z singapurense
5b. Bracts tightly appressed, bracteoles well-developed (25-30 mm) 6
Singapore native Zingiberales I.
161
6a. Spike ovoid, bracts bronze to red-purple at anthesis, labelliim pale yellow
ornamented by fine dark purple lines and tinged in the centre Z. elatius
6b. Spike narrowly fusiform, bracts orange to rose-pink at anthesis, labellum plain
pale yellow 7
7a. Leafy shoots 2-3 m tall, inflorescence to 110 cm long (peduncle to 65 cm long,
spike to 44 cm long) Z. petiolatum
7b. Leafy shoots 1-2 m tall, inflorescence to 56 cm long (peduncle to 40 cm long,
spike to 1 6 cm long) Z. aurantiacum
The genus Zingiber in Singapore
In Singapore only two Zingiber species are cun'ently considered native: Zingiber
griffithii Baker and Zingiber puberulum Ridl. Both are easy to distinguish from
Zingiber singapurense even when sterile (Fig. 4). Zingiber griffithii has leafy shoots
0.5-0. 8 m tall, with 5-8(-10) leaves per shoot, leaf sheaths almost glabrous at base,
with sparse long silky appressed hair at apical part, lamina elliptic, thin, bright green,
slightly plicate, c. 1 2-25 x 5-8 cm, with long silky appressed hairs abaxially. Zingiber
puberulum is a more robust species with leafy shoots up to 2 m tall, with up to 23 leaves
per shoot, leaf sheaths puberulous to pubescent, lamina elliptic, slightly leathei'y, mid
to dark green, sometimes with dull silvery finish, almost flat (plication negligible), c.
20-35 X 5-8 cm, adaxially puberulous. Zingiber singapurense is a delicate plant, not
exceeding 0.8 m in height (though its arching leafy shoots might be up to 1.5 m long),
with 18-26 leaves per shoot, lamina narrowly ovate, slightly leathery, dark glossy
green, flat (not plicate) 15-22 x 2-A cm, glabrous adaxially, lighter green and almost
glabrous abaxially (except occasional space hairs near the midrib and the basal part
near pulvinus). The above measurements given are from living material in Singapore
populations.
The habit of Zingiber singapurense blends very well with other understorey
vegetation and at first glance is reminiscent of young palm fronds or even some
terrestrial orchids (e.g. Bromhaedia finlaysoniana (Lindl.) Miq.). Unless it is flowering
it is fairly hard to spot this rare species, which is likely the reason why it has been
overlooked until now.
Tliree varieties of Zingiber puberulum are currently recognised (Holttum, 1950;
Theilade 1998), Z. puberulum var. puberulum, Z. puberulum var. chryseum (Ridl.)
Holttum and Z. puberulum var. ovoideum (Ridl.) Holttum), of which two are based on
Singapore material. Zingiber puberulum war. puberulum was described from material
collected in Bukit Timah Nature Reserve (Singapore) where it is still present. All
extant Shigapore material of this species belongs to this variety. Zingiber puberulum
var. chryseum was described in 1908 by Ridley (as Z. chryseum) from the woods in the
Stagmount area (Singapore). The forest in this area was cleared shortly thereafter and
no further material of this taxon has ever been collected. It is presumed, therefore, to
162
Gard. Bull. Singapore 66(2) 2014
Fig. 3. Zingiber elatius (Ridl.) Theilade, photographed at Penang Hill (type locality). A. Habit
and detail of ligule (inset). B. Inflorescence and detail of flower (inset). Zingiber aurantiacum
(Holttum) Theilade, photographed at Fraser’s Hill (type locality). C. Habit and detail of ligule
(inset). D. Inflorescence and detail of flower (inset). (Photos: Jana Leong-Skomickova)
Singapore native Zingiberales I.
163
Fig. 4. Zingiber griffithii Baker, photographed in Bukit Timah Nature Reserve, Singapore
{SNG-127). A. Habit and detail of ligule (inset). B. Infloreseence shortly after flowering.
Zingiber puberulutm var. puberulum Ridl., photographed in Bukit Timah Nature Reserve,
Singapore (type loeality; SNG-91). C. Habit and detail of ligule (inset). D. Infloreseence (image
v
rotated 90° counter clock-wise). (Photos: Jana Leong-Skomickova)
164
Gard. Bull. Singapore 66(2) 2014
have gone extinct in Singapore. Zingiber ptiberulum van ovoideum was described by
Ridley from material collected in Pahang and so far this taxon has been only reported
from Peninsular Malaysia (Kelantan, Perak and Pahang). Its record for Singapore by
Chong et al. (2009) is an error and should be corrected to Z. puberulum war. ptiberulum.
Zingiber ottensii Valeton (previously recorded as Z zerumbet e.g. by Keng et
al., 1998; Chong et ah, 2009) has been reported growing in gardens and waste ground
(Keng et al., 1998), and later listed as casual (a sub-category of an exotic species) in a
Checklist ofthe total vascular plantflora of Singapore (Chong et al., 2009), As with many
other widely utilised gingers, the exact origin of Zingiber ottensii is unclear, though
it is clearly native to SE Asia. This species is distributed in the evergreen Malesian
region, compared to the closely related Zingiber zerumbet (L.) Sm. which is mostly
confined to South Asia and monsoonal parts of Southeast Asia (Indochinese region).
The two species are indeed very similar when sterile and practically indiscernible in
herbarium material even if an inflorescence is present unless good notes on the flowers
are provided. It is, therefore, not surprising that these are often confused. Zingiber
ottensii when flowering can be readily distinguished from Z. zenimbet by the rusty
purple-brown colour of the bracts (compared to bright green bracts of Z. zerumbet)^
and by the labellum and stammodes, which are tinged by pale purple to violet and have
pale yellow blotches (compared to the pure cream-yellowish labellum and staminodes
of Z. zerumbet) (Fig. 5). So far we have seen numerous populations of Zingiber ottensii
freely growing in several locations in Singapore but have not observed any populations
of Zingiber zerumbet.
Key to native and naturalised Zingiber species in Singapore
la. Spike ovoid, bracts rusty purple brown at anthesis, peduncle equal or more than
double the length of spike (> 15 cm long), labellum and staminodes tinged by
light purple to violet, with light yellow blotches Z. ottensii
lb. Spike fusifonn, bracts pink, orange or red at anthesis, peduncle as long or shorter
than spike (< 15 cm), labellum and staminodes unifonnly pale yellow 2
2a. Plant less than 1.5 m tall, bracteole less than 6 mm or absent 3
2b. Plant over 1.5 m tall, bracteole more than 10 mm Z. puberulum
3a. Lamina elliptic, bright green, thin, weakly plicate, beneath with long silky
appressed hair, bracteoles absent Z. griffithii
3b. Lamina narrowly ovate, dark green, slightly leathery, flat (not plicate), glossy,
glabrous beneath, bracteoles reduced (2-7 mm long) Z. singapurense
ACKNOWLEDGEMENTS. We sincerely thank Matti Niissalo (National University of
Singapore) and Boo Chih Min (Uvaria Tide). Each brought to our attention one additional
population of Zingiber singapurense. We thank the National Parks Board, Singapore, for the
Singapore native Zingiberales I.
165
Fig. 5. Zingiber zerumbet (L.) Sm., photographed in southern Vietnam. A. Infloreseenee.
B. Flower (front view). Zingiber ottensii Valeton, photographed in Singapore, outskirts of
Bukit Timah Nature Reserve. C. Infloreseenee. D. Flower (front view). (Photos: Jana Leong-
Skomickova)
166
Gard. Bull. Singapore 66(2) 2014
generous funding of our project ‘Enhancement of Biodiversity Research for Gingers’ (January
2011 - March 2012) through the Urban Ecology RF programme. This enabled extensive
surveys across Singapore leading to this and other exciting discoveries, as well as mass
propagation of material of various native Zingiberales for re-introductions. Our sincere thanks
are due to nmnerous NParks statT and volunteers, who continuously supported us in the field
during plant collections and regular monitoring, in particular: Mishak Sunari (Sungei Buloh
Wetland Reserve), Derek Liew, Chung Yi Fei, William Ng and Ngon Soon Kong (all Central
Catchment Nature Reserve), Tran Hun Dang (ex staff of Singapore Botanic Gardens), Sunia
Teo (Singapore Botanic Gai'dens), Michael Leong (volunteer, Singapore Botanic Gardens).
We also thank Adelle Wang (Bukit Batok Nature Park), Desmond Lee (Sungei Buloh Wetland
Reserve) for their active approach in re-introduction programmes, and Koh Teng Seah and Dr
Gillian Khew (both Singapore Botanic Gardens) for supporting in vitiv propagation of native
gingers at Singapore Botanic Gardens. We also thank Dr Axel Poulsen and an anonymous
reviewer for critical comments and suggested improvements to this manuscript. Dr David
Middleton, Editor of Gardens’ Bulletin Singapore, for language improvements and navigating
the paper through the review process, and the curators of KEP and SING herbaria for allowing
us to examme relevant herbarium material.
References
Bastmeijer, J.D. & Kiew, R. (2001). A new Oyptocoryne hybrid (Araceae) from the Bukit
Timah Nature Reserve, Singapore. Gard. Bull. Singapore 53: 9-17.
Chong, K.Y., Tan, H.T.W. & Corlett, R.T. (2009). A checklist of the total vascidar plant flora
of Singapore: native, naturalised and cultivated species. Singapore: Raffles Museum of
Biodiversity research, National Llniversity of Singapore.
Chong, K.Y., Tan, H.T.W. & Corlett, R.T. (2011). A summary of the total vascular plant flora
of Singapore. In: Wong, K.M., Leong-Skomickova, J., Lee, S. & Low, YW. (eds)
Proceedings of the 8th Flora Malesiana Symposium. Gard. Bull. Singapore 63: 197-204.
Cowley, J. & Theilade, I, (1995). 267. Zingiber sulphureum, Zingiberaceae. Bot. Mag. 12:
73-77.
Davison, G.W.H. (2008). The Red List categories. In: Davison, G.W.H., Ng, P.K.L. & Ho, H.C.
(eds) The Singapore Red Data Book: Threatened plants and animals of Singapore, 2nd
ed. Pp. 1-4. Singapore: The Nature Society (Singapore).
Holttum, R.E. (1950). The Zingiberaceae of the Malay Peninsula. Gard. Bull. Singapore 13:1-
249.
lUCN (2012). lUCN Red List Categories and Criteria: Version 3.1. Second edition. Switzerland,
Gland and UK, Cambridge: lUCN.
Kishor, R. & Leong-Skomickova, J. (2013). Zingiber kangleipakense (Zingiberaceae): Anew
species from Manipur, India. Gard. Bull. Singapore 65(1): 39-46.
Keng, H., Chin, S.C. & Tan H.T.W (1998). The Concise Flora of Singapore, volume II:
Monocotyledons. Singapore University Press.
Lim, C.K. (2003). Zingiber aurantiacum (Holtt.) Theilade, Z petiolatum (Holtt.) Theilade and
two related new taxa from Peninsular Malaysia and Thailand. Folia Malaysiana 4(2):
65-76.
Low, Y.W., Leong, P.K.F., Tee, S.P., Rajesh Singh, Tay, M.L.C. & Wong, K.M. (2014).
Margaritaria (Phyllanthaceae), a new generic record for the Singapore flora. Gard. Bull.
Singapore 66(1): 47-56.
Singapore native Zingiberales I.
167
Ng, P.K.L., Corlett, R.T. & Tan, H.T.W. (2011). Singapore Biodiversity. An Encyclopaedia
of the Natural Environment and Sustainable Development. Editions Didier Millet in
association with the Raffles Museum of Biodiversity Research.
Rodda, M., Tran, H.D., Chiew, P.T., Liew, D. & Leong-Skomickova, J. (2012). The rediscovery
of Dischidia hirsuta (Apocynaceae, Asclepiadoideae) in Singapore. Card. Bull.
Singapore 64(2): 293—299.
Sugumaran, M. & Wong, K.M. (2014). Studies in Malesian Gentianaceae, VI. A revision of
Utania in the Malay Peninsula with two new species. PI. Ecol Evol. 147(2): 213-223.
Theerakulpisut, R, Triboun, P. Mahakham, W, Maensiri, D., Khampila, J. & Chantharanothai,
R (2012). Phylogeny of the genus Zingiber (Zingiberaceae) based on nuclear ITS
sequence data, 5w//. 67: 1-7.
Theilade, I. (1998 [‘1996’]). Revision of the genus Zingiber in Peninsular Malaysia. Gard.
Bull. Singapore 48: 207-236.
Wu, T.L. & Larsen, K. (2000). Zingiberaceae. In: Wu, Z.Y. & Raven, RH. (eds) Flora of China
24: 333-346. Beijing: Science Press.
Yao, T.L. (2013). Nine new species of Thottea (Aristolochiaceae) in Peninsular Malaysia and
Singapore, with two taxa m Peninsular Malaysia redefined and a taxon lectotypified.
Blurnea 58(3): 245—262.
Yee, A.T.K., Corlett, R.T., Liew, S.C. & Tan, H.T.W. (2011). The vegetation of Singapore -
an updated map. In: Wong, K.M., Leong-Skomickova, J., Lee, S. & Low, Y.W. (eds)
Proceedings of the 8th Flora Malesiana Symposium. Gard. Bull. Singapore 63(1 &2):
205-212.
Yeoh, Y.S., Yeo, C.K., Ang, W.F. & Low, Y.W. (2013). Marsdenia maingayi (Apocynaceae,
Asclepiadoideae), a rare rainforest woody climber rediscovered in Singapore. Gard.
Bull. Singapore 65: 241-249.
Gardens’ Bulletin Singapore 66(2): 169-171. 2014
169
Three new combinations in Haplopteris
(Pteridaceae subfam. Vittarioideae)
S. Lindsay^ & C.W. Chen^
'Gardens by the Bay, 18 Marina Gardens Drive,
Singapore 018953
Stuart. 1 i ndsay @garden sby th ebay.com .sg
^Department of Molecular and Cellular Biology,
National Tsing Hua University,
Hsinchu, Taiwan 30013
ABSTRACT. Three new combinations are made here: Haplopteris alternans (Copel.) S. Linds.
& C.W.Chen, Haplopteris humblotii (Hieron.) S.Linds. & C.W.Chen, and Haplopteris
dareicarpa (Hook.) S.Linds. & C.W.Chen. Lectotypes for Vittaria alternans Copel. and Vittaria
humblotii Hieron. are also designated here.
Keywords. Adiantaceae, Haplopteris, Monogramrna, Pteridaceae subfam. Vittarioideae,
Vittaria, Vittariaceae, Vittarioid
Introduction
In 1998 Crane published a revised circumscription of the genera of the fern family
Vittariaceae (now Adiantaceae p.p. or Pteridaceae subfam. Vittarioideae p.p.) based on
a phylogenetic analysis of rbcL gene sequences and supported by micro-morphological
characters. He proposed a number of generic changes and many new combinations but
the most significant change for Asia was the loss of the genus Vittaria Sm. (Vittaria s.s.
was redefined as a small genus of approximately 6 species restricted to the Neotropics
and Africa) and the resurrection of the genus Haplopteris C.Presl to accommodate
the many Asian species formerly placed in Vittaria. Most of the necessary new
combinations in Haplopteris have already been made (see Crane, 1998; Zhang, 2003;
Lindsay, 2010) but a few species such as Vittaria alternans Copel. from the Pliilippines
and Vittaria humblotii Hieron. from the Comoros and Madagascar appear to have been
overlooked. Both of these species have soral paraphyses with funnelfonn apical cells,
a character that they share with all existing species of Haplopteris but which is absent
from Vittaria s.s. In view of this, two new combinations in Haplopteris need to be
made.
Although Crane (1998) clarified the phylogenetic relationship of most genera of
vittarioid ferns his work did not include the two small genera Rheopteris Alston and
Monogramrna Comm, ex Schkuhr s.l. A decade later, Ruhfel et al. (2008) published
the results of another phylogenetic analysis of rhcL sequence data that included these
genera. This analysis included Rheopteris cheesmaniae Alston, the sole member and
170
Gard. Bull. Singapore 66(2) 2014
therefore the type of the genus Rheopteris, as well as four species of Monogramma
s.l {Monogramma dareicarpa Hook., M. paradoxa (Fee) Bedd. (as Vaginularia
angustissima (Brack.) Mett.), M, trichoidea J.Sm. ex Hook, and M. acrocarpa
(Holttuni) D.L.Jones) but not, unfortunately, Monogiwnma graminea (Poir.) Schkiihr,
the type of the genus. Despite the absence of Monogramma graminea, this study
clearly showed that M. paradoxa, M. trichoidea and M. acrocarpa form a strongly
supported monophyletic group that is sister to Rheopteris cheesmaniae and that this
clade is sister to the rest of the vittarioid ferns. Monogramma dareicarpa, conversely,
is nested within Haplopteris. Moreover, Monogramma dareicarpa has bilateral spores
and soral paraphyses with fimnelform apical cells, characters that are shared with most
species of Haplopteris, whereas M. paradoxa, M. trichoidea and M. acrocarpa have
trilete spores and soral paraphyses with unspecialised apical cells. On the basis of
these and other differences, and moiphological differences to M. graminea, Ruhfel et
al. (2008) argued that the genus Vaginidaria Fee, a genus rarely recognized in recent
literature but of which Vaginularia {Monogramma) trichoidea is the type, should be
separated from Monogramma s.s. Ruhfel et al. (2008) also discussed the implications
of Monogramma dareicarpa being nested within Haplopteris but were reluctant to
formally transfer it to Haplopteris without knowing the placement of M. graminea.
The main concern was that \ f Monogramma graminea were to be nested \n Haplopteris
then, under the principle of priority, and without conservation, the genus Haplopteris
would become a synonym of Monogramma and the combination in Haplopteris would
be pointless. There are also a number of key morphological differences between
Monogramma graminea and M. dareicarpa that make assumptions of their relationship
to each other problematic. Very recently the phylogenetic position of Monogramma
graminea has been investigated using sequence data from four chloroplast genes (C.W.
Chen, unpublished). Its precise placement in the phylogeny of vittarioid ferns is to be
published separately after broader sampling and further analysis but it is already clear
that Monogramma graminea is (1) not closely related to M. dareicarpa, (2) is not
nested in Haplopteris and (3) is not closely related to any of the sampled Vaginularia
species. These facts support our decision to also transfer Monogramma dareicarpa to
Haplopteris.
New combinations
Haplopteris alternans (Copel.) S.Linds. & C.W. Chen, comb. nov. — Vittaria alternans
Copel., Philipp. J. Sci. 1 (Suppl. 2): 157 (1906). TYPE: Philippines, Mindanao,
Zamboanga, San Ramon, 2600 feet (800 m), 2 May 1905, In tree top in upland forest,
E.B. Copeland 1767 (lectotype MICH [MICHl 191098], designated here; isolectotype
BM (fragment) [BM000008869]).
Haplopteris humblotii (Hieron.) S.Linds. & C.W.Chen, comb. nov. — Vittaria
humblotii Hieron., Bot. Jahrb. Syst. 53(3-5): 427 (1915). TYPE: Comoros, Island of
Three new combinations in Haplopteris
171
Anjouan, January 1887, L. Humblot 1526 (lectotype P [P00149188], designated here;
isolectotypes P (x3) [P00149187, P00149189, P00149190]).
Haplopteris dareicarpa (Hook.) S.Linds. & C.W.Chen, comb. nov. — Monogramma
dareicarpa Hook., Sp. Fil. 5: 121, t.288A (1864). — Pleurofossa dareicarpa (Hook.)
Nakai ex H.Ito., J. Jap. Bot. 12: 408 (1936). TYPE: Borneo, Labiian [Pulau Labuan,
Sabah, Malaysia], s.d.. On the trunks of trees, E.S. Barber [IMotley] s.n. (holotype K)
Note. A letter (dated 16 November 1853) in the archives at Kew from Edmund Scott
Barber, Resident Director of the Eastern Archipelago Company at Labuan, to William
Jackson Hooker reveals that Barber purchased a collection of more than 400 botanical
specimens fi*om James Motley (who he replaced at the Eastern Archipelago Company)
in 1 853 and, soon after, sent these to Hooker. Barber only arrived in Labuan from
England in 1 853 and died the following year so it is possible that many of the Bomean
specimens that are attributed to Barber, including the type of M. dareicarpa, should
be attributed to Motley instead. Motley lived in Labuan from 1849 to 1853 (and then
elsewhere in the region until his death in 1859) and is reported to have been a keen
botanist and collector of natural history specimens.
References
Crane, E.H. (1998 [1997]). A revised eircumseription of the genera of the fern family
Vittariaeeae. Syst. Bot. 22(3): 509-517.
Lindsay, S. (2010). New combinations in Haplopteris (Adiantaceae) for the Flora of Peninsular
Malaysia. Card. Bull. Singapore 62(1): 119-120.
Ruhfel, B., Lindsay, S. & Davis, C.C. (2008). Phylogenetic placement of Rheopteris and the
polyphyly of Monogramma (Pteridaceae s.l.): Evidence from rbcL sequence data. Syst.
Bot. 33(1): 37-43.
Zhang, X-C. (2003). New combinations in Haplopteris (Pteridoph 5 ^a: Vittariaeeae). Ann. Bot.
Fennici 40(6): 459-461.
Gardens’ Bulletin Singapore 66(2): 173-188. 2014
173
Puccinia psidii (Pucciniaceae - Eucalyptus Rust, Guava
Rust, Myrtle Rust) - a threat to biodiversity
in the Indo-Pacific region
R.O. Makinson'’^ & BJ. Conn^
'National Herbarium of New South Wales, Mrs Macquaries Road,
Sydney NSW 2000 Australia
bob.makinson@rbgsyd.nsw.gov.au
barry.conn@rbgsyd.nsw.gov.au
^Australian Network for Plant Conservation Inc.,
GPO Box 1 777, Canberra ACT 2601 Australia
ABSTRACT. A biotype of the South American fimgal rust pathogen Puccinia psidii Winter has
become widely naturalised along the east coast of AusPulia since early 2010, reaching North
Queensland in 2012. This pathogen is known globally as Eucalyptus Rust or Guava Rust, and
in Australia as Myrtle Rust. It is pathogenic on a wide range of plants in the Myrtaceae. This
pathogen constitutes a major threat to myrtaceous plants in natural habitats and in production
systems in moister ai'eas of Australia, and potentially on a wide scale in the Malesian region.
The risk of spread to the Malesian region, from Australia or other sources, is high, and once
naturalised in Myrtaceae-rich biomes, eradication is unlikely.
Keywords. Eucal 5 dpus Rust, Guava Rust, Myrtle Rust, Puccinia psidii, Pucciniaceae
Introduction
In late April 2010, a disease was identified on cultivated specimens of native plants in
the family Myrtaceae near Sydney, initially only in a few commercial plant nurseries.
The disease pathogen was rapidly identified as an exotic rust fungus, either Puccinia
psidii Winter (Pucciniaceae, order Pucciniales), or a close relative. Puccinia psidii,
which is of South American origin and is internationally known as Eucalyptus Rust
or Guava Rust, had been regarded for many years as a major potential threat to native
Australian and Malesian species of the Myrtaceae if it ever arrived in the region. In
Australia, the pathogen was initially regarded as the uredinial morpho-species Uredo
rangelii J.A.Simpson, K.Thomas & C.A.Grgurinovic, a taxon putatively narrowly
distinct from P. psidii (Simpson et al., 2006). The name Uredo rangelii was widely
used for the Australian outbreak during 2010-2011. Subsequent genetic studies have
concluded that tliis pathogen is a biotype of Puccinia psidii (Carnegie & Cooper,
2011; Carnegie et al., 2010; Pegg et al., 2013, 2014) and this is now the scientifically
recognised name for the pathogen that is present in Australia, with Myrtle Rust being
the vernacular name that is in wide use. Much of the electronic and printed material
on the Australian incursion dating from 2010-2012 used the name 'Uredo range IW .
174
Gard. Bull. Singapore 66(2) 2014
The change of preferred scientific name does not invalidate that literature in most
other respects, although host-lists and range reports from that period are now greatly
outdated. In this paper, we use the tenn Myrtle Rust for the variant (biotype) present in
Australia, and the composite term Eucalypt/Guava Rust for the total species {Puccinia
psidii) of which it is part. Note that Myrtle Rust is not to be confused with ‘Myrtle
Wilt’; a fungal disease of Myrtle Beech {Nothofagus cunninghamii (Hook.) Oerst.,
Nothofagaceae) caused by the pathogen Chalara australis J.Walker & G.A.Kile
(Smith, 2004).
Myrtle Rust was initially known only from nursery and cut-flower production
facilities, and eradication was attempted (Cannon, 2011; Carnegie & Cooper, 2011),
with infected stock being traced and destroyed. However, by October 2010 there were
many reports along the New South Wales coast, most resulting from human movement
of infected plant material, and the first reports of naturalisation. In late December
2010 the pathogen was deemed to be ineradicable and the emergency response was
discontinued. In 2011, the disease naturalised along the east coast of Australia from
southern New South Wales to south-eastern Queensland, and has since spread north
as far as Cooktown in northern Queensland. Outbreaks have occurred in the southern
State of Victoria since late 2011, mainly in and around Melbourne; the disease is now
regarded as endemic in that State but, to June 2014, occurrences there remain confined
to plants in open cultivation and to plant production and distribution facilities and it has
not naturalised in natural vegetation (D. Smith, Victorian Department of Environment
and Primary Industries, pers, comm. June 2014).
This paper briefly reviews the biology, ecology and global spread of this
pathogen, its host range, field recognition and symptoms, effects on individual hosts,
implications for conservation, and the available management guidelines, information
and training resources.
Potential scale of the problem
In the early stages of contingency planning for Puccinia psidii a national scoping
document (Commonwealth of Australia, 2006) rated it as “one of the most serious
threats to Australian production forests and natural ecosystems ... It has a potential to
cause direct mortality in the estimated 1 0% of all Australian native forest plant species
(and the great majority of dominant species) that belong to the family Myrtaceae, and
with indirect effects that may include habitat loss for native fauna and flora, retarded
regeneration and recruitment of younger trees and successional species, greater
impact of fire, and abiotic effects as a result of canopy decline including erosion,
reduced water quality, reduced water retention in soil and vegetation and potentially
large losses through lost production to the forestry industry.” In the later National
Contuigency Plans (Plant Health Australia, 2008, 2009), the pathogen was assessed
as having a high potential for entry into Australia, a high establishment potential, a
high-to-extreme spread potential, a high environmental impact, and a high-to-extreme
economic impact. It was stated that “The climate in Indonesia and Papua New Guinea
Eucaljq^tus Rust in Indo-Pacific
175
is expected to be very hospitable for P. psidiP (Plant Health Australia, 2009, p. 73
& Fig. 4), based on the predictive map of Magarey et al. (2007) which used broad
climatic factors (temperature and rainfall) to indicate that most of New Guinea and
Indonesia are potentially suitable for the establishment of Puccinia psidii.
Most pre- and post-amval impact assessment and plaiming for Australia has
been by primary industries organisations (Cannon, 2011; Nursery & Garden Industry
Australia, 2012; Plant Health Australia, 2009; Tommerup et ah, 2003), with an
emphasis on horticultural and forestry systems. Planning and assessment for natural
biodiversity has lagged behind badly, although given some attention by Glen et al.
(2007), NSW Office of Environment and Heritage (201 1); NSW Scientific Committee
(2011); Pegg et al. (2013). Plamiing and impact assessment has been hampered by
inadequate taxonomic and epidemiological knowledge for the several to many variants
of Puccinia psidii, and by an unavoidable early reliance on mainly South American
data regarding host range and climatic tolerances.
Global spread of Eucalypt/Guava Rust {Puccinia psidii)
The presumed origin and centre of diversity of Puccinia psidii is in Brazil and Uruguay
(South America). In this region, the effects are usually mild on native host species (Glen
et al., 2007; Perez et al., 2011). The pathogen came to notice through its sometimes
severe effects on crop species of Myrtaceae that are not native to the region, including
species of Psidium L. (Guava) and Eucalyptus L’Her. sens. lat.
In the first half of the twentieth century Puccinia psidii spread to the Caribbean,
where the allspice industry {Pimenta dioica (L.) Merr.) was severely damaged. In
1977, a variant arrived in Florida (U.S.A.), where among other hosts it attacked the
Australian species Melaleuca quinquenervia (Cav.) S.T.Blake, a naturalised woody
weed of the Everglades.
In 2005 a variant arrived in Hawaii, and within a year had spread to nearly all
the Hawaiian Islands, where it is now known as Ohia Rust. Its effects have been severe
on one native Hawaiian species, the already endangered Eugenia koolauensis O.Deg.,
with “a very high proportion” of plants killed (Cannon et al., 2010, p. 47). It has
affected some naturalised non-Hawaiian Myrtaceae, especially Rose Apple, Syzygium
jambos (L.) Alston (Loope, 2010).
In 2007 Eucalypt/Guava Rust was reported from Japan (Kawanisihi et al., 2009),
where it may have since been eradicated, in 20 1 1 from Hainan, China (Zhuang & Wei,
2011), and in 2013 Ifom New Caledonia (Giblin, 2013) and South Africa (Roux et al.,
2013).
In Indonesia, Hardiyanto & Tridasa (2000) report a 'Puccinia sp.’ rust disease
on sapling eucalypts in 1998 during forestry trials in Kalimantan and Sumatra. The
host eucalypts were hybrids developed in China but involvmg parent stock from
Brazil. The disease seems to have died out at these sites and has apparently not been
confiiiTied as Puccinia psidii.
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Gard. Bull. Singapore 66(2) 2014
The mode of arrival in Australia is not known, and the plant nursery sites where
it was first reported seem not to have been arrival sites.
On the Myrtaceae
The Myrtaceae has its centre of diversity in the Australasian region, where it is
structurally and floristically dominant in many ecosystems, and ecologically significant
in many more.
Australia has about 2,253 native species of the family Myrtaceae in 88 genera
(B. Lepschi, Australian National Herbarium, pers. comm., Aug. 2010). Myrtaceae
comprise about 10% of Australia’s native flora. Roughly half the Australian species
occur m the climatic zones likely to be most conducive to Eucalypt/Guava Rust.
The Malesian region has a less speciose and less generically diverse myitaceous
flora than Australia, but the family is still very well -represented. Comprehensive data
are lacking, but exemplar numbers are available for some areas. New Guinea and the
Solomon Islands have about 300 species in 28 genera (Hoft, 1992). A similar number
of myitaceous species, in about 30 genera, occur in Indonesia (Craven et al., 2003),
with 26 species, in 6 genera in Central Kalimantan (Campbell, 1999). 216 tree species,
in 9 genera are recorded for Peninsular Malaysia (Kochummen, 1978), and more than
205 species, in 14 genera in Sabah and Sarawak (Malaysia) (Ashton, 2011).
The Indo-Pacific region as a whole is very depauperate in rust fimgi that
parasitise Myrtaceae (Walker, 1 983). In this region, nearly all rusts on Myrtaceae are,
or are likely to be, of introduced (non-native) origin. Only two rusts on myitaceous
plants are known to occur naturally in Australia and both are rare (Simpson et al.,
2006). Walker (1983, pp. 116 & 117) reports “two unconfirmed records from India of
Melampsora ... on two introduced Australian species of Eucalyptus''. As a result of
the near-total isolation from myitaceous rusts, the Myrtaceae of the region are thought
to be epidemiologically ‘naive’ to these pathogens, with no co-evolved defences.
However, resistance of various sorts may exist through autochthonous traits.
Biology and ecology of Puccinia psidii
Taxonomic and pathotypic knowledge of Puccinia psidii is still incomplete, but there
appear to be several biotypes, all of which infect only plants of the Myrtaceae. These
biotypes may differ in host preferences, severity on different hosts, and environmental
tolerances.
The life cycle of Puccinia psidii is described in Glen et al. (2007), and, with
some new information, in Morin et al. (2014). In brief, Puccinia psidii is functionally
autoecious (it can complete its life cycle on a single host plant), and hemicyclic (no
spemiogonia and aecia ever observed - Glen et al. (2007), but see Morin et al. (2014)
for qualifications). The asexual uredinial life cycle can be completed an indefinite
number of times and produces infective urediniospores in great numbers (e.g. Uchida
Eucaljq^tus Rust in Indo-Pacific
111
& Loope, 2009). The iiredinial cycle may take as little as 10 days from infection to
sporulation under suitable conditions. Urediniospores disperse readily by wind, by
animal vectors, and through human movement of infected material. At least some
variants, including the one in Australia, also produce sexual teliospores, which are
fewer, larger, and less mobile, but may allow adaptive variation (Glen et al., 2007),
although their functionality in the life cycle of the Australian variant is uncertain
(Morin et al., 2014).
Both the geimination and pustule growth stages of the pathogen require
susceptible host species, suitable tissues (typically new leaves and very young stems),
and favourable ambient conditions. The spore germination stage requires wet leaf
surfaces at night within a suitable temperature range. Both germination and pustule
fonnation require moderate temperatures, usually cited (e.g. Glen et al., 2007) as
13-22 (-2 5 )°C based on South American data. However, there are indications that,
in the Myrtle Rust variant, pustules may still form at lower temperatures (albeit at
lower frequency), with early winter overnight minima as low as of 8°C (G. Guymer,
Queensland Herbarium, pers, comm. 2011) or even 2-5°C (D. Smith, Victorian
Department of Primary Industries, pers. comm. June 2012).
Tn Hawaii, Puccinia psidii “has been found statewide ... attacking Myrtaceae
from near sea level to elevations of about 1200 m in areas with rainfall ranging from
750-5000 mm” (Loope, 2010, pp 1 & 6). In Australia, occurrences of sporulating
Myrtle Rust from higher elevations have been recorded at 600 m (Canberra, latitude
35°S); 1 000 m (Mount Tomah, NSW, 33”S); 700 m (Toowoomba, Queensland, 27.5°S),
and c. 1 000 m (Mt Hypipamee National Park, Queensland, 17.4°S).
Urediniospores are very light and very mobile in the air column. There is a
noticeably high incidence of infections starting on the lower surface of leaves, only
later penetrating to the upper surface. Nursery inspection strategies need to take
account of this.
Some South American data (Zauza et al., 2010b) suggest a gradient of infection
frequency with height in plantations of eucalypt saplings, attributed to higher spore
loads lower in the air column, and to the upper canopy being drier at night compared to
lower leaves and hence less conducive to spore germination. Several South American
studies suggest lower levels of infection in mature eucalypts, even on seasonal new
growth, as opposed to juvenile (sapling stage) plants.
The hyphae of genninating spores penetrate the plant though the waxy cuticle,
not through the stomates as with some other rusts (K. Old in Cannon et al., 2010; Hunt,
1968). Initial symptoms can appear within 5-7 days of infection. The whole uredinial
reproductive cycle can occur in 10-12 days in warmer months, and depending on
the initial number of infection sites a massive outbreak of pustules with a very high
new spore load can develop very quickly. The cycle slows or stalls in cold months at
temperate latitudes.
Spore longevity is generally cited (often following Glen et al., 2007) as 90
days for reduction to low levels of viability, but probably varies greatly with ambient
conditions. Lower longevity occurs at temperatures >30‘'C. Salustiano et al. (2008)
report the survival levels of spores under various storage conditions, noting that spores
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Gard. Bull. Singapore 66(2) 2014
in deep-freeze and liquid nitrogen storage maintained significant levels of viability at
150 days. Although timber and pulp production facilities were likely to receive spores
in “very low numbers, the adverse environmental conditions encountered in these areas
and during overseas transport do not favour spore survival. Thus, the risk of spread
of this pathogen into new areas in the absence of infected host plants is considered
extremely low to inexistent” (Lana et al., 2012, p. 1). This finding is inconsistent with
the report of Grgurinovic et al. (2006) that viable spores were detected on a wood
shipment to Australia from South America. Further study of longevity is required
to enable general and specific risk assessments, including for seed-banks handling
potentially contaminated seed-lots. Long-distance transmission to new areas via spore
loads on human clothing and equipment (e.g. hats, rucksacks, tents) is distinctly
possible (Tommerup et ah, 2003).
Phylogeny and taxonomy of the pathogen
Separate lines of DNA investigation confirm that Pticcinia psidii is phylogenetically
misplaced in genus Puccinia and the family Pucciniaceae (van der Merwe et al. 2008;
Pegg et ah, 2013; Tan et ah, 2014; E. Liew et al. unpublished data, pers. comm. June
2014). Its affinities within the order Pucciniales are as yet unresolved.
Host species taxonomic range
Since the arrival of Puccinia psidii in Australia, over 300 native taxa (>10% of the
total Austi*alian Myrtaceae) have been recorded as hosts of the pathogen, either in field
situations (wild or cultivated), or under laboratory test conditions (for Australian State
host lists, see below; Carnegie & Lidbetter, 2011; Morin et ah, 2012). The number
of hosts is expected to increase further. Based on field studies, almost 45% of the
Australian myrtaceous genera (51 out of 88, see Appendix) have one or more species
known to be susceptible, affecting almost all Myrtaceae tribes represented in Australia
(14 tribes out of 17).
Separate but overlapping lists of Australian hosts have been maintained by the
Australian States in which Myrtle Rust currently oceurs, and are accessible via links
from Plant ffealth Australia (2012); these lists are however not fully up to date as
at June 2014. Consolidated and up to date global and Australian host lists are now
(October 2014) available on-line (Giblin & Carnegie 2014a, b).
There is no clear phylogenetic pattern of host susceptibility, with wide variation
in apparent susceptibility of species within genera, and sometimes within species.
Susceptibility may vary with host genetic factors (resistance of a species or individual
plant), host physiological and stress factors, growth stage, and environmental factors
including host density, local spore load, and suitability of conditions for spore
germination and pustule growth. Severity ratings are as yet tentative and, to date, are
Eucaljq^tus Rust in Indo-Pacific
179
only being gathered and published by the Queensland Department of Agriculture,
Fisheries and Forestry.
Syzygium jamhos is very susceptible in both Australia and Hawaii, with young
leaves, flower buds and petioles all prone to conspicuous levels of pustule formation
(Fig. 1). This makes it an excellent indicator species, but also a prime source of local
spore load if not dealt with promptly.
It is likely that several of the Austiulian native host species will undergo severe
decline over time as a result of repeated cycles of infection by Puccinia psidii, and that
some species not cuiTently regarded as threatened with extinction will become so -
this is already evident for the hitherto common mesic shrubs Rhodomyrtus psidioides
(G.Don) Benth. and Rhodamnia rubescejis (Benth.) Miq. (A. Camegie & G. Pegg,
unpublished data, pers. comm. March-May 2014). Some of the more susceptible
species are important floristic elements in native ecological communities, and their
decline may have a pronounced effect on those communities. The high susceptibility
of new growth on A4elaleuca quinquenervia (Cav.) S.T.Blake, M leucadendm (L.) L.
and M. viridiflora Sol. ex Gaertn. is of great concern, given the extensive floodplain
and riparian associations in which they occur in northern Australia.
The potential long-term effects on eucalypts are unclear. Some 80 eucalypt
taxa, not counting hybrids, are recorded as susceptible - 69 taxa in Eucalyptus, eight
in Cofymbia, and three in Angophora (A. Camegie, G. Pegg, F. Giblin unpublished
data, pers. comm. March 2014; Morin et al., 2012). However, in Eucalyptus itself
a majority of these records are from experimental laboratory trials, including on
non-eastern Australian species, and only a minority have infection confinued in the
natural environment. No eucalypts are regarded as highly susceptible at this point,
but there is a lack of observations for nearly all species on the critical life stages of
seedlings and coppice growth (e.g. post-fire epiconnic shoots). Infection of cotyledons
in cultivated seedlings of Eucalyptus planchoniana F.Muell. resulted in death of the
whole batch (Pegg et al., 2013). This suggests a potential for high mortality during the
seedling phase of susceptible eucalypts subjected to high mst spore loads in conditions
favourable for spore germination.
Field recognition of Puccinia psidii
Puccinia psidii is not known to utilise as host any plants other than members of the
Myrtaceae (including Heteropyxidaceae). Suspected occurrences of Puccinia psidii
should be reported as soon as possible to national and local biosecurity authorities and
lead conservation agencies. The initial report should be accompanied, if possible, by
digital photographs of the symptoms and of the host plant, the latter showing enough
features to enable identification. Australian diagnostic services may be able to assist
(biosecurity@industry.nsw.gov.au or www.daff.qld.gov.au/4790_20842.htm - for
phytosanitary reasons, send photographs only, not samples). Laboratory confirmation
of the pathogen may be needed.
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For images of the various stages of Puccinia psidii infection on a variety of
host species and tissues, refer to Plant Health Australia (2012) for links to websites
maintained by the Australian State governments of New South Wales, Queensland and
Victoria. Early symptoms of infection, small purplish or brown-grey lesions, are not
obvious and are difficult to distinguish from other minor damage; they typically occur
on new stem and leaf tissue, but also on soft fruits and parts of inflorescences in some
hosts. Leaf infections often start on the lower surface, penetrating to the upper surface
in 2-4 days. Within a few more days sporulating pustules develop.
The sporulation (pustule) stage is the easiest to recognise. By the time this stage
is obvious, spores are ready to disperse - handling and disposal of plants at this stage
can easily spread the spores unless strict protocols are followed. The most prolific
form is the uredinial pustule, typically bright yellow or yellow-orange (Fig. 1, 2). Less
common are light- to mid-brown telial pustules, producing sexual teliospores.
After the spore masses have dispersed, the effects on the plant may remain
obvious (e.g. dead shoots, defoliated new stems, distorted new stem structure and
overall habit) but are not easy to attribute definitely to being a Puccinia psidii infection
in the absence of pustules.
Effects of infection
The effects of severe infection may include a loss of new seasonal or regenerative
growth, some loss of photosynthetic capability, loss of some or all reproductive capacity
if fiowers or fruits (soft-fruited species) are infected or if flowering can only occur
on new growth, exhaustion of growth potential, and death or retardation of seedlings
and juveniles. Whole plant death is known in plantations of some eucalypt and other
species in South America after repeated infection cycles. Whole-plant mortality in
Australia has, so far, mainly been seen in some seedlings, nursery stock, and new bush
regeneration plantings, but is now increasingly seen in naturally occurring plants of the
highly susceptible Rhodamnia rubescens and Rhodomyrtus psidioides.
Repeated infection cycles on highly susceptible hosts may cause decline in
population size or density, with effects on pollination systems and gene flow. Animals
dependent on myrtaceous species may be affected. Regional or total extinction of very
susceptible species is possible, as are stRictural and ecological changes in vegetation
communities and faunal assemblages.
Climatic & geographic prediction modelling
Predictive modelling has been limited, and largely dependent on matching South
American climatic, microclimatic and epidemiological data (often from plantation
conditions and of uncertain Puccinia psidii biotypes) to Australian and Melanesian
bioclimatic profiles.
Eucal3q3tus Rust in Indo-Pacific
181
Fig. 1. Uredinial pustules of Puccinia psidii on new growth of eultivated Syzygium jambos
(Rose Apple), Brisbane, Queensland, Sept. 2011. (Photo: R.O. Makinson)
Booth et al. (2000) produced a preliminary assessment of high risk areas for
the Northern Territory, Queensland and New South Wales. A revised continental
risk-predictive map by Booth & Jovanovic appeared in Glen et al. (2007) (as a pers.
comm.; refer Booth et al., 2000, and Booth & Jovanovic, 2012 for the methodology
used to develop map) and was reproduced in Plant Health Australia (2009). The latter
publication also featured an alternative map (p. 14) by R. Magarey that includes
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Gard. Bull. Singapore 66(2) 2014
Fig. 2. Uredinial pustules of Puccinia psidii on hypanthium and ovary summit of Rhodomyrtus
psidioides (Native Guava), Wamberal Nature Reserve, New South Wales, Dee. 2010. (Photo:
R.O. Makinson)
Australia, New Guinea and most of Indonesia, taken from a global risk analysis (based
on distributions in South and Central America) later published as Magarey et al.
(2007). The Malesian zones on this map are assessed as having a very high likelihood
of permanent Eucalypt/Guava Rust establishment. Elith et al. (2012) developed a
model for investigating the differences in distribution of artificially grouped locations
in Australia that they deemed represented different strains of the rust.
The most recent modelling, of relative climate suitability for Puccinia psidii
at global and Australian scales, is that of Kriticos et al. (2013). This shows most of
Malesia and parts of mainland South-east Asia as having moderately to highly suitable
climatic parameters for the permanent establishment of Puccinia psidii, assuming
favourable local climatic regimes and the presence of susceptible host species.
Impact assessment and management guides for production systems
The most comprehensive management guide available for the horticultural sector is
the Australian Nursery Industry Myrtle Rust Management Plan (Nursery & Garden
Industry Australia, 2012). This contains detailed monitoring, biosanitation, and
fungicide treatment protocols for production and wholesale/retail ‘greenlife’ facilities.
This Plan is also applicable to forest nurseries and to some plantation situations.
Eucaljq^tus Rust in Indo-Pacific
183
Short management guidelines for various situations in the Australian context,
including domestic and rural properties, and for industries based on the myrtaceous
oils and foodstuffs, are available via the Queensland and New South Wales primary
industries department websites (Plant Health Australia, 2012). The New South Wales
Road Transport Authority Biodiversity Guidelines provide model procedures for
reducing the risk of spread by utilities providers and contractors (NSW Roads and
Maritime Services, 2011).
A broad assessment of the potential impact of this disease on the Australian
commercial forestry sector is provided by Cannon (2011), who comments that
long-term effects will depend partly on whether other strains of Eucalypt/Guava
Rust are introduced. Camegie et al. (2010) and Pegg et al. (2014) provide details of
methods used for rapid screening of (eucalypt) forestry species for susceptibility.
Some recent research projects funded by the Australian Government have a bearing
on resistance in the production context (publications pending by; B. Thumma et al.,
CSIRO Plant Industry, resistance genes in eucalypts; C. Kulheim et ah, Australian
National University, genetic markers for resistance in non-eucalypt Myrtaceae, and
metabonomic responses to infection; K. Sandhu & R.F. Park, University of Sydney,
genetic basis of pathogenicity; M. Horwood, University of Sydney, chemical control).
A considerable literature, mostly from South America, exists for the eucalypt
plantation forestry sector, including research on pathotypes, resistance genetics, and
resistance breeding - key recent papers, with more or less extensive reference lists,
are da Silva et al. (2014), Gra9a et al. (2011), Mamani et al. (2010), Martins et al.
(2011) and Zauza et al. (2010a). A consolidated interim global bibliography for R
psidii (Makinson 2014) is now available on-line, although still incomplete for South
America.
Prescriptions for the assessment of impact and the management of Puccinia
psidii in forestry and in the wild are at an early stage of development in Australia. The
impacts of the pathogen, and the (limited) options for its management, are gradually
becoming apparent. There is a time-limited opportunity, especially in those susceptible
areas not yet infected by Myrtle Rust, to gather baseline data and to establish long-
term monitoring. In the absence of any likely biological control, effective management
in extensive Myrtaceae-rich systems of a multi-vector disease with such a wide host
range is inherently problematic.
Impact assessment and management guides for wild biodiversity
Impact assessment for Australian native {in situ) biodiversity has been slow in
developing, and only sparse, mostly qualitative data are available. One early
assessment with some national applicability, is for the state of New South Wales (NSW
Scientific Committee 2011). Work in progress by G. Pegg (Queensland Department of
Agriculture, Forestry and Fisheries) and A. Camegie (New South Wales Department
of Primary Industries) on a few wild species represents the main organised research
on the impacts on natural species to date. They report (pers. comm. May 2014) severe
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Gard. Bull. Singapore 66(2) 2014
declines in seasonal growth, reproductive capacity and recruitment in the mesic shrubs
Rhodomyrtus psidioides and Rhodamnia rubescens.
There are no ‘good’ management solutions for Myrtle Rust infection in natural
vegetation. Fungicidal treatments are rarely possible on the scale needed and have
unwanted effects on other biodiversity. Physical removal of susceptible species is
usually ineffective and too expensive except in small areas, and may be incompatible
with conservation management objectives. However, such a strategy may have value
(e.g. along high-use tracks) in reducing spore load on visitors likely to then ti'avel to
rust-free zones. Fire is unlikely to be an effective control given the potential for re-
infection and the preference of the rust for new growth.
The likely behaviour of Myrtle Rust in relation to local floristic composition
and recmitment, microclimate, disturbance (especially fire), and regeneration is not
known. In climatically marginal areas, or where highly susceptible host species are
lacking (or die out), infections in natural vegetation may be transitory or seasonal.
Some frrst-generation bushland policy/management documents are now
available, including NSW Office of Environment and Heritage (2011). Summary
adviee for minimising arrival risk and impact on natural plant communities, including
hygiene protocols, is available as brochures and webpages (refer the Australian
government primary industries sites of affected States: Plant Health Australia 2012).
Conclusion
Puccinia psidii poses a signifrcant threat to the Myrtaceae flora of the Malesian and
Australasian regions, and to at least some ecological communities. It is understood
that some level of regional infonnation exchange has been developed through primary
industries and quarantine agencies. It is highly desirable that this be formalised and
expanded to include environmental organisations (govenunent and non-government),
and other stakeholders.
ACKNOWLEDGEMENTS. We thank Angus Carnegie (New South Wales Department of
Primary Industries), Fiona Giblin (University of the Sunshine Coast), .lonathan Lidbetter
(New South Wales Department of Primary Industries), Geoff Pegg (Queensland Department
of Agriculture, Forestry and Fisheries), David Smith (Victorian Department of Environment
and Primary Industries), Gordon Guymer (Queensland Herbarium), and Edward Liew (Royal
Botanic Gardens and Domain Trust, Sydney) for providing unpublished infonnation that has
been incorporated into this paper. Saw Leng Guan (Forest Research Institute Malaysia) kindly
assisted us to develop an estimate of the species of Myrtaceae in Malaysia.
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Appendix. Host genera of Myrtle Rust (Puccinia psidii) in Australia, to 22 June 2014.
Legend: Bold = one or more species of genus infected in wild or in cultivation; underlined =
infectivity in one or more species of genus by artificial inoculation under laboratory conditions
only (Morin et ah, 2012); * = genera enthely exotic to Australia; f = genera native to Malesia.
Generic recognition follows Wilson et al. (2005) note that some other published lists follow
differing lists of recognised genera. Unpublished data courtesy of J. Lidbetter and A. Carnegie
(New South Wales Department of Primary Industries), F. Giblin (University of the Sunshine
Coast), and G. Pegg (Queensland Department of Agriculture, Forestry and Fisheries). Note that
susceptibility of a species in a genus does not necessarily imply susceptibility of all congeners.
Acmena, Acmenosperma^ Agonis^ Allosvncarpia. Anetholea, Angophora^ 'fAsteromyrtus,
Austromyrtus, Backhousia, f Baeckea, Barongia, Beaufortia. Callistemon, Calothamnus,
Calvtrix ., Chamelaucium, Choricarpia^ Corymbia, Darwinia^ '\Decaspenniim, '\Eucalyptus,
■fEugenia^ fCossia, *Heteropyxis, Homoranthus, Hypocalymma^ KunzecL Lenwebbia,
"^Leptospennum, "fLindsayomyrtus, Lithomyrtus, Lophomyrtus^ fLophostemon,
^Melaleuca, ^Metrosideros, Mitrantia, "^^Myrciaria, "^Myrtus, ^Osbornia, Pilidiostigma^
* Pimenta . Plinia, *f Psidium . Re^elia. ^Rhodamnia, fRhodomyrtus, Ristantia, Sphaerantia,
Stockwellia^ Syncarpia, 'fSyzygiiim^ Thi'vptomene . Tristania^ t Tristanionsis . *Ugni,
•fUromyrtus, Verticordia . Waterhousea, Xanthostemon.
Gardens’ Bulletin Singapore 66(2): 189-194. 2014
189
A new species of Billolivia (Gesneriaceae) from Vietnam
DJ. Middleton^, J, Leong-Skomickova^ & Q.B. Nguyen^
'Herbarium, Singapore Botanic Gardens,
National Parks Board, 1 Cluny Road, Singapore 259569
david_middleton@nparks ,gov. sg
-Vietnam National Museum of Nature,
Vietnam Academy of Science and Teclmology,
1 8 Hoang Quoc Viet Street, Cau Giay, Hanoi, Vietnam
ABSTRACT. The new species Billolivia moelleri D.J.Middleton fromNui Chua National Park
in Niiih Thuan Province in Vietnam is described. A provisional lUCN conservation assessment
and a new key to the species of Billolivia are provided.
Keywords. Billolivia, Gesneriaceae, new species, Vietnam
Introduction
The genus Billolivia D.J.Middleton, containing five species new to science, was only
recently described (Middleton et al., 2014). These five species are known from a
relatively small area in the southern Annamite mountains in the provinces of Binli
\
Phuoc and Lam Dong in Vietnam. Although one of the collections was made in 1933,
all of the other material from which these species were described was collected within
the last 20 years and most of it within the last 10 years. However, the collecting density
in the region remains fairly low. As the known species of Billolivia are all quite locally
endemic the likelihood that new species in the genus will be discovered as forested
areas in the region are better explored is high. A collection made in November 2013
in Nui Chua National Park in Ninli Thuan province has already proven to be a new
species in the genus and is here described. A provisional lUCN conservation assessment
following the guidelines in lUCN (2012) is suggested.
Billolivia moelleri D.J.Middleton sp. nov.
Similar to Billolivia violacea D.J.Middleton & H.J.Atkins in the violet corolla lobe
tips and the presence of a calyx tube but differing in the leaf margins being more
coarsely dentate, in the dark purple lines in the corolla, and in the corolla being smaller
overall. It also shows similarities to Billolivia vietnamensis D.J.Middleton & Luu in
the overall corolla size and in the purple lines in the corolla but that species has the
calyx divided to the base and the corolla lobes are white.
TYPE: Vietnam, Ninh Thuan Prov., Ninh Hai Dist., VTnh Hai Commune, Nui Chua
National Park, trek from Kien Kien village to Nui Chua peak, 758 m, 11°44’27.4”N
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109°07’41.9”E, 2 November 2013, Jana Leong-Skornickovd, Nguyen Quoc Binh,
Aung Thame & Edward Ong JLS-2623 (holotype SING [including flowers and fruits
in spirit]; isotypes E, K, MO, P, VNMN). (Fig. 1 , 2)
Caulescent herb to 30 cm tall; stems with long brown hairs to 4.5 mm long. Leaves
alternate, intemodes up to 2.9 cm long but generally around 1 cm long; petioles 6.5-
1 1 .2 cm long, densely covered in a mixture of long brown spreading hairs to 6 mm long
and shorter appressed brown hairs; lamina elliptic to obovate, 7.6-14.5 x 3. 1-6.2 cm,
2.1-3 times as long as wide, base cuneate to somewhat truncate, apex short acuminate,
margin dentate to crenate, more coarsely so towards apex, secondary venation 8-12
veins on each side of midrib with weaker intersecondaries between, tertiary venation
alternate percurrent, adaxial lamina with sparse long hairs throughout, margin ciliate,
abaxial lamina with shorter hairs to 1 mm long between the veins and both short
and long hairs to 4.5 mru long on midrib and venation. Inflorescences subsessile,
c. 8-flowered; bracts at base, narrowly triangular, 0.6-1 x 1.5-1 .7 mm, apex acute;
pedicels 28-33 mm long, densely long pubescent. Calyx 14-15 mm long, of a short
tube and 5 lobes, somewhat 2-lipped with the three upper lobes close and parallel and
the two lower lobes diverging, outside densely covered in long eglandular hairs to 5
mm long, inside glabrous except for few minute papillae; tube 4-5.5 mm long; lobes
deltoid, apices acute, 9-10 x l .5-2 mm. Corolla c. 25 mm long, composed of a nan'ow
tube which slightly curves downwards in lower half and slightly flares towards mouth,
and a 2-lipped limb, white with purple lines from inside tube and onto lobes, ventral
surface of throat and tube with 2 yellow stripes in line with lobe sinuses, upper halves
of lobes violet; tube c. 15 mm long; upper lip 2-lobed, c. 11 mm long, sinus between
lobes c. 4 mm, lobes 4x6 mm; lower lip 3-lobed, c. 13 mm long, lobes obovate, apices
rounded, lateral lobes c. 9 x 7 mm, medial lobe c. 10x9 mm; corolla outside glabrous
at base of tube, with short hairs on tube above this and then with short and long hairs on
upper part of tube and outside of lobes, inside glabrous in tube and with short glandular
hairs on inside of lobes. Stamens inserted at c. 9 mm from corolla base; filaments c.
7 mm long, narrow at base, widening around middle and curved and twisting at this
point, with short stalked glands in upper half; 2 fertile anthers c. 2 x 1.8 mm, thecae
± parallel, 2 anthers fused by tips; lateral staminodes c, 2 mm long; medial staminode
c. 1 miu long. Disc aimular, 0.2 mm high. Ovary c. 5 mm long, glabrous at base of
ovary and glandular pubescent in upper half; style c. 1 1 mm long, glandular pubescent
throughout with hairs of widely vai 7 ing lengths; stigma 2-lobed, lobes c. 1. 1 mm long.
Fruit an ellipsoid berry, pale green or with a pink tinge, 10-12.5 mm long, 4-6 mm
wide, pubescent distally. Seeds numerous, unappendaged, c. 0.6 x 0.4 mm.
Distribution. Vietnam. Only known from the type collection from Nui Chiia National
Park.
Habitat. Understorey of lower montane broadleaved evergreen forest, usually growing
on steep slopes and in moist gulleys.
New Billolivia from Vietnam
191
Fig. 1. Billolivia moelleri D.J.Middleton. A. Habit. B. Flowers, ^xom Leong-Skornickovd et al.
JLS-2623. (Photos: Jana Leong-Skornickova)
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Gard. Bull. Singapore 66(2) 2014
Fig. 2. Billolivia moelleri D. J.Middleton. A. Detail of flower (side view). B. Detail of flower
(front view). C. Stem with fruits and leaves, adaxial view. D. Detail of fruits. From Leong-
Skornickovd et al. JLS-2623. (Photos: Jana Leong- Skornickova)
New Billolivia from Vietnam
193
Etymology. The species is named in honour of Dr Michael Mdller of the Royal Botanic
Garden Edinburgh for his contribution to our understanding of the Gesneriaceae in
Asia, including as a co-author on the paper in which the genus BiUolivia was first
described.
Provisional lUCN Conservation Assessment. Vulnerable VU D2. This species is only
known from the type locality where the plant is not commonly encountered. It is
difficult to estimate the population size or the exact Area of Occupancy but it is likely
to be around 20 km^ as the altitude at which the species is found restricts the available
vegetation in the National Park. Within the Park itself though there are no immediately
discernible tlireats to qualify it as threatened under the B category.
Key to Billolivia species
la. Calyx divided into 5 lobes almost to base; corolla lobes white or white with
purple veining at base with tips white, corolla tube 8-18 mm long (unknown in B.
poilaneiy, leaf laminas minutely dentate or crenate, often appearing entire 2
lb. Calyx connate into a tube at base for at least 4 mm, margin 5-lobed; corolla lobes
white at base, tips brightly coloured pink, red or violet, corolla tube 15-28 mm
long; leaf laminas minutely or conspicuously dentate 4
2a. Calyx lobes > 4 mm wide B. poilanei
2b. Calyx lobes < 2.5 mm wide 3
3a. Leaf abaxially with hairs only on veins or with only occasional hairs on lamina;
corolla 12-15 mm long; unfertilised ovary glabrous B. minutiflora
3b. Leaf abaxially with pubescence tlu'oughout; corolla 18-25 mm long; unfertilised
ovary pubescent at apex B. vietnamensis
4a. Corolla tube c. 15 mm long, inside of throat white with purple lines in tube and
onto lobes B. moelleri
4b. Corolla tube 20-28 mm long, inside of throat white with no coloured lines 5
5a. Calyx fused into a tube for 7-9 mm; corolla lobe tips pink or red; leaf lamina
margins coarsely dentate; petioles 9-18 cm long B. longipetiolata
5b. Calyx flised into a tube for 4-6 mm; corolla lobe tips violet; leaf margins minutely
dentate or appearing entire; petioles 6-12.5 cm long B. violacea
ACKNOWLEDGEMENTS. The research of the first and second authors is funded by the
National Parks Board, Singapore. We would like to thank the People’s Committee and the
Agriculture and Rural Development Department of Ninh Thuan Province, Mr Dang Kim
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> r
Cuong, the Director of Nui Chua NP, and Mr Tran Van Tiep, the Head of the Department of
Science (Nui Chiia NP), for issuing permits and for giving various logistical and personnel
support for our fieldwork. We thank Assoc. Prof Dr Nguyen Trung Minh, Dr Vu Van Lien
and Prof Lim Dam Cu from the Vietnam National Museum of Nature for their support of our
collaborative research. We thank Mr Aung Thame and Mr Edward Ong from Singapore Botanic
Gardens for their assistance in the field.
References
lUCN (2012). lUCN Red List Categories and Criteria: Version 3.1. Second edition. Switzerland,
Gland and UK, Cambridge: lUCN.
Middleton, D.J., Atkins, H., Luu, H.T., Nishii, K. & Moller, M. (2014). Billolivia, a new genus
of Gesneriaceae from Vietnam with five new species. Phytotaxa 161: 241-269.
Gardens’ Bulletin Singapore 66(2): 195-205. 2014
195
A new genus of Gesneriaceae in China and the transfer of
Briggsia species to other genera
M. Moller' ^ W.-H. Chen'-^, Y.-M. Shui', H. Atkins^ & D.J. Middleton'*
^Key Laboratoiy of Plant Diversity and Biogeography of East Asia,
Kunming Institute of Botany, Chinese Academy of Sciences,
132 Lanhei Road, Kunming 650201, Yunnan, China
^Royal Botanic Garden Edinburgh, 20A Inverleith Row,
Edinburgh EH3 5LR. Scotland, UK
m.moeller@rbge.ac.uk.
^University of the Chinese Academy of Sciences,
Beijing 100049, China
"^Herbarium, Singapore Botanic Gardens,
National Parks Board, 1 Cluny Road, Singapore 259569
ABSTRACT. Since the transfer of several species of Briggsia to Oreocharis, including the type
species Briggsia longifolia, the remaining 16 species of Briggsia have been in taxonomic limbo.
We address this unsatisfactory situation by transferring 1 0 further species into Oreocharis on
morphological grounds and by raising a new genus, with two species, based on previously
published molecular data and their morphological distinction from other genera. This leaves
only four species for which, at present, no satisfactoiy solution is available but, for pragmatic
reasons until further research can be done, we place them in Loxostigma to which they are
moiphologically most similar and in which one already has a combination. Several names are
lectotyprfied.
Keywords. Briggsia, Didymocarpinae, Gesneriaceae, Glabrella, lectotypification, Loxostigma,
new combinations, Oreocharis
Introduction
Species placed in the now synonyniised genus Briggsia Craib in the Gesneriaceae,
subtribe Didymocarpinae (Weber et ah, 2013), have a complex taxonomic history,
being moved in and out of a number of genera as taxonomic concepts changed, e.g.
Roettlera Vahl, Didymocarpus Wall., Chirita Buch.-Ham. ex D.Don, Didissandra
C.B.Clarke dind Loxostigma C.B. Clarke. These numerous changes did not necessarily
indicate a progressively better understanding of these taxa but rather the difficulties
of placing species displaying conflicting characteristics in genera that were narrowly
defined primarily on floral characteristics.
The genus Briggsia was established by Craib (1919a) with three new species.
In a separate paper, but which was published in the same issue (Craib, 1919b), he
transfeiTed 1 1 species from Didissandra to Briggsia and defined Briggsia as having
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species with a large, distinctly bilabiate ventricose corolla with gradually inarching
filaments and four anthers cohering in pairs, irrespective of other characters such as
growth habit as both caulescent and acaulescent species were included.
Of the tliree new species in Craib (1919a), Briggsia longifolia Craib was later
designated as the lectotype (Burtt, 1954). Craib (1919b) described an additional
variety along with the transfers from Didissandm. His concept of Briggsia included
two truly caulescent species, i.e., B. amabilis (Diels) C.B.Clarke (later synonymised
into B. kurzii (C.B.Clarke) W.E.Evans) and B. cavaleriei (H.Lev. & Vaniot) Craib
(later transferred to Loxostigma as L. cavaleriei (H.Lev. & Vaniot) B.L. Burtt). It also
contained Briggsia longipes Craib that can produce short stems up to six cm, with
the remaining 11 species being acaulescent and rosette-forming. Chun (1946) later
added two more rosette species, Burtt (1955) described a further rosette species, and
Burtt (1958) transfeiTed the rosette species Briggsia rosthornii (Diels) B.L.Burtt from
Didissandra. In Pan (1988) one species was synonymised (Briggsia fritschii = B.
mihieri) and six new species and three varieties were added to the genus, including two
further caulescent species, B. longicaulis W.T.Wang & K.Y.Pan and B. dongxingensis
Chun ex K.Y.Pan, with stems 20 to 60 cm long. In Wang et al. (1990), Pan reduced
Briggsia crenulata Hand.-Mazz. to a variety of B. rosthornii (B. rosthornii var.
crenulata (Hand.-Mazz.) K.Y.Pan). Lastly, a further truly caulescent species, Briggsia
damingshanensis L.Wu & B.Pan was recently described (Wu et al., 2012), bringing
the total to 22 species and four varieties. Briggsia dulongensis H.Li is an invalid name
without a Latin description (Li, 1993).
The advent of molecular phylogenetic methods has enabled us to examine
problematic species’ placement with new suites of characters. These techniques have
already proven useful for delimiting other problematic Gesneriaceae genera (Weber et
al., 2011a, 201 lb, 201 Ic; Middleton & Moller, 2012; Middleton et al., 2013) and have
also led to the descriptions of new genera (Wei et al., 2010; Middleton & Moller, 2012;
Middleton et al., 2014, submitted). Moller et al. (2011b) used the results of molecular
phylogenetic studies to greatly enlarge the genus Oreocharis Benth. to include all
species of nine other genera (Ancylostemon Craib, Bournea Oliv., Dayaoshania
W.T.Wang, Deinocheilos W.T.Wang, Isometrum Craib, Opithandra B.L.Burtt,.
Paraisometiwn W.T.Wang, Thamnocharis W.T.Wang and Tremacron Craib), and
part of Briggsia. In Moller et al. (2011b) five species and four varieties of Briggsia
were transfeiTed to Oreocharis, including the type species B. longifolia, leaving the
remaining species in a taxonomic limbo. On the basis of additional molecular studies
on the enlarged genus Oreocharis, a ftirther species, Briggsia speciosa Craib, is in the
process of being transferred to Oreocharis by Chen et al. (2014). All Briggsia species
transferred to Oreocharis thus far have been rosette-forming plants. This leaves
16 species unplaced, including the caulescent species. The molecular phylogenetic
studies conducted so far indicate that these caulescent species are not closely related to
the acaulescent species now placed in Oreocharis (Moller et al., 2011a). Furthermore,
the truly caulescent Briggsia dongxingensis and B. kurzii do not form a clade. They
Anew Gesneriaceae genus and the transfer oiBriggsia species
197
appear in two different positions in the phylogenetic trees so far published and are in
unsupported positions with respect to clades containing Petrocosmea Oliv., Loxostigma
and Pseudochirita W.T.Wang species, inteimixed with Raphiocarpus Chun species.
For Briggsia damingshanensis no molecular data are available, but the species differs
only marginally from B. dongxingensis from the same region in Guangxi province
(Wu et al, 2012). The (indirect) link of the tmly caulescent species to Loxostigma is
interesting since in the Flora of China (Wang et al., 1 998), their generic placement in
Briggsia was questioned by Weitzman and Skog who suggested that B. dongxingensis
and B. longicaulis probably belong to Loxostigma. Vitek et al. (2000) placed Briggsia
kurzii in Loxostigma but acknowledged that this has not been followed by other
authors. However, though phylogenetically close, a case for the placement of these
truly caulescent species in Loxostigma is not easily made since Loxostigma is closely
related to Pseudochirita. Loxostigma Pseudochirita share the presence of rhizomes
and mainly differ in the former having four stamens and appendaged seeds, the latter
two stamens and unappendaged seeds. Rhizomes, diandry and appendaged seeds are
unknown in the truly caulescent Briggsia species. Because of these morphological
differences and their uncertain phylogenetic position a taxonomic placement of these
species is, at present, difficult to predict. However, given their current nomenclaturally
unacceptable position we include them in Loxostigma where Briggsia kurzii already
has a combination. We are well aware, however, that more work is needed here and
their status may change in the future.
Briggsia longipes and B. mihieri form a strongly supported sister clade. Their
phylogenetic position is isolated from the other former Briggsia species and all
other didymocaipoid genera (Mdller et al., 2011a) and a new genus is necessary to
satisfactorily place these species. The two species can be clearly differentiated from
other species fonnerly placed in Briggsia by their glabrous petiole and leaf blade (only
puberulous when young), combined with their short, 5-6 cm, stems. The remaining
ten unplaced Briggsia species are truly stemless and possess a dense pubescence on
the petioles and adaxial and abaxial leaf surfaces. These are characteristics they share
with most members of the expanded Oreocharis and the transfers are, therefore, made
here. This is preferred to the alternative of withholding their transfer until leaf material
for DNA analysis eventually becomes available. This might be years since some
species are rare and have not recently been collected and thus the herbarium material
is also unsuitable for DNA extraction. The traditional taxonomic approach using
moiphological data was undertaken in the case of the dismantling of Chirita (Weber et
al., 2011a) for those species that were not included in the molecular analyses, and for
the recently described Somrania D.J.Middleton which was established on the basis of
moiphology alone (Middleton & Triboun, 2012; see also Puglisi, 2014).
We have designated a number of lectotypes below. In each case we have
chosen the most complete specimen showing the range of characters from amongst
the available candidates. An appendix is attached listing all Briggsia names and their
current status.
198
Gard. Bull. Singapore 66(2) 2014
Taxonomic Treatments
Glabrella Mich.Moller & W.H.Chen, gen. nov.
Differs from other Chinese genera of Gesneriaceae by the combination of indistinct
short stem, 5-6 cm long; glabrous petiole and leaf blade, leaf base cuneate or narrowly
peltate; fertile stamens 4, cohering in pairs at apex; capsule straight in relation to
the pedicel, not twisted; and seeds unappendaged. TYPE: Glabrella mihieri (Franch.)
Mich.Moller & W.H.Chen
Plants stemless or stems to 5-6 cm tall, glabrous. Leaves basal or crowded at apex of
short stem; petiole glabrous; leaf blade narrowly obovate to elliptic, glabrous, apex
rounded to acute; lateral veins 3 to 5 on each side of midrib. Cymes few-flowered;
peduncle (5-)8-23 cm; bracts 2, linear to narrowly triangular or lanceolate, 1-4 x
0.5- 1.5 mm, glabrous, margin entire. Calyx segments lanceolate to narrowly ovate to
narrowly triangular, margin entire. Corolla blue-purple or pale purple to pale yellow,
inside usually spotted, (3.2-)4-6 cm, outside glabrous to sparsely glandular pubescent
or pubemlent, inside pubemlent; tube 2.1-4 x l.l-L8(-2.6) cm; adaxial lip 5-9 mm,
lobes semiorbicular, 3-7 x 6-9 mm, apex rounded; abaxial lip 1-1.4 cm, lobes oblong
to semiorbicular, 6-7 x 4-8 mm, apex obtuse to rounded. Adaxial stamens adnate to
corolla 8-9 mm above base, abaxial ones adnate to corolla 8-1.2 cm above base, 1.2-
1.7 cm long; filaments glabrous or sparsely glandular pubescent; anthers ovoid, thecae
not confluent; staminode 0.8-1 mm. Pistil 2.5-3 cm; ovary pubescent; style 0.7-2 mm,
glabrous to sparsely pubescent. Capsule 3.4-7 cm, glabrescent, straight in relation to
the pedicel, not twisted. Seeds numerous, unappendaged, FI. Sept-Oct, fr. Nov-Dec.
Distribution. Two species, endemic to China.
Etymology. After the distinctly hairless leaves.
Key to Glabrella
la. Leaf blade base cuneate, margin crenate-serrate; calyx segments 4-7 x 1.5-3
mm; peduncle glabrous to glabrescent G. mihieri
lb. Leaf blade base peltate or rounded to nearly cuneate, margin entire to shallowly
serrate; calyx segments 8-11 x 2-5 mm; peduncle sparsely brownish villous,
rarely glabrescent G. longipes
Glabrella longipes (Hemsl. ex Oliv.) Mich.Moller & W.H.Chen, comb. nov. —
Didissandra longipes Hemsl. ex Oliv., Hooker’s Icon. PI. 24: pi. 2379 (1895); Hemsl.,
Bull. Misc. Inform. Kew 1895: 114 (1895). — Briggsia longipes (Hemsl. ex Oliv.)
Craib, Notes Roy. Bot. Gard. Edinburgh 11: 262 (1919). TYPE: China, Yunnan
province, Mengtze, Dec 1843, W. Hancock 50 (holotype K! [K000858088]).
Anew Gesneriaceae genus and the transfer oiBriggsia species
199
Distribution. China: Chongqing (Hechuan Xian), Guangxi (Longlin Xian, Tianlin
Xian), SE Yunnan.
Glabrella mihieri (Franch.) Mich.Moller & W.H.Chen, comb. nov. — Didissandra
mihieri Franch., Bull. Mens. Soc. Linn. Paris 1: 450 (1885). — Didymocarpus
mihieri (Franch.) H.Lev., Compt. Rend. Assoc. Frang. 34: 427 (1906), nom. inval. —
Briggsia mihieri (Franch.) Craib, Notes Roy. Bot. Gard. Edinburgh 11: 262 (1919).
TYPE: China, Kouy-Tcheou (Guizhou) province, 1858, RH. Ferny s.n. (holotype P!
[P03511310]).
Didissandra fritschii H.Lev. & Vaniot, Compt. Rend. Assoc. Frang. 34: 425 (1906).
— Didymocarpus fritschii (H.Lev, & Vaniot) H.Lev., Compt. Rend. Assoc. Fran9. 34:
428 (1906), nom. inval. — Briggsia fritschii (H.Lev. & Vaniot) Craib, Notes Roy. Bot.
Gard. Edinburgh 11: 262 (1919). TYPE: China, Guizhou, Tsin-gay, Montagues du
Lion, 24 Oct 1898, Laborde & Bodinier 2464 (holotype E! [E00387546]).
Distribution. China: Guangxi (Longlin Xian), Guizhou, S Sichuan.
Loxostigma C.B. Clarke
Loxostigma damingshanensis (L.Wu & B.Pan) Mich.Moller & H.Atkins, comb. nov.
— Briggsia damingshanensis L.Wu & B.Pan, Ann. Bot. Fenn. 49(1-2): 79 (2012).
TYPE: China, Guangxi, Nanning, Damingshan Natural Reserve, 23 28 ’N, 108 25 ’E,
alt. 1250 m, 5 Aug 2010, Lei Wu & Rihong Jiang D0320 (holotype IBK!; isotypes
IBK!, PE).
Distribution. China: Guangxi (Nanning).
Loxostigma dongxingensis (Chun ex K.Y.Pan) Mich.Moller & Y.M.Shui, comb. nov.
— Briggsia dongxingensis Chun ex K.Y.Pan, Acta Phytotax. Sin. 26: 451 (1988).
TYPE: China, Guangxi, Dongxing, Shiwandashan, X.R.Liang 70078 (holotype IBSC
[0649548], image seen; isotype A [A00353708], image seen).
Distribution. China: Guangxi (Dongxing Xian); N Vietnam.
Loxostigma kurzii (C.B. Clarke) B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 34:
104 (1975). — Didymocarpus kurzii C.B. Clarke, Commelyn. Cyrtandr. Bengal, t. 66
(1874). — Chirita Ixurzii (C.B.Clarke) C.B. Clarke, J. Linn. Soc. 15: 145 (1876). -
Roettlera kurzii (C.B.Clarke) Kuntze, Rev. Gen. PI. 2: 476 (1891). — Briggsia kurzii
(C.B.Clarke) W.E.Evans, Notes Roy. Bot. Gard. Edinburgh 16: 133 (1928). TYPE:
India, Sikkim, Kursiong, Kurz s.n. (holotype CAL n.v.).
200
Gard. Bull. Singapore 66(2) 2014
Didissandra amabilis Diels, Notes Roy. Bot. Gard. Edinburgh 5 : 224 (1912). — Briggsia
amabHis (Diels) Craib, Notes Roy. Bot. Gard. Edinburgh 1 1 : 263 (191 9). TYPE: China,
Yunnan, Tali Range [now Dali Range], Forrest 2689 (lectotype E! [E00387561],
designated by Vitek et al. (1998); isolectotypes E! [E00387559, E00387560]).
Briggsia amabilis var. taliensis Craib, Notes Roy. Bot. Gard. Edinburgh 1 1 : 263 (1919).
TYPE: China, Yunnan, eastern flank of the Tali Range [now Dali Range], Lat. 25° 40’
N, alt. 9,000-1 0,000 ft., fl. Jul-Aug 1 906, G. Forrest 4385 (lectotype E! [E00387582],
designated here; isolectotype IBSC [1BSC0004824], image seen).
Distribution. China: SW Sichuan, NW Yunnan; NE India: Sikkim; Nepal; Bhutan;
Myanmar [Burma]
Loxostigma longicaule (W.T.Wang & K.Y.Pan) Mich.Moller & Y.M.Shui, comb. nov.
— Briggsia longicaulis W.T.Wang & K.Y.Pan, Acta Phytotax, Sin. 26: 450 (1988).
TYPE: China, Sichuan, alt. 2500 m, 24 Aug 1959, Exped. PL Econ. Liangshan 5836
(holotype PE [PE00030680], image seen; isotype PE [PE00030681], image seen).
Distribution. China: Sichuan (Butuo Xian, Dechang Xian, Kangding Xian).
Oreocharis Benth.
Oreocharis acutiloba (K.Y.Pan) Mich.Moller & W.H.Chen, comb. nov. — Briggsia
acutiloba K.Y.Pan, Acta Phytotax. Sin. 26: 455 (1988). TYPE: China, Yunnan, Yuxi,
alt. 2250 m, 23 Sept 1958, S.K. Wu 61 (holotype KUN! [KUN484366]).
Distribution. China: Yunnan (Yuxi Xian).
Oreocharis agnesiae (Forrest ex W.W.Sm.) Mich.Moller & W.H.Chen, comb. nov.
— Didissandra agnesiae Forrest ex W.W.Sm., Notes Roy. Bot. Gard. Edinburgh 8:
334 (1915). — Briggsia agnesiae (Forrest ex W.W.Sm.) Craib, Notes Roy. Bot. Gard.
Edinburgh 1 1 : 263 (191 9). TYPE: China, Yunnan, Mountains of the Yung Peh, Lat. 26°
40’ N, alt. 9,000-10,000 ft, fl. Jul 1914, G. Forrest 12,829 (lectotype E! [E00135143],
designated here; isolectotypes E! [E00135142, E00135144], K! [K000858909], IBSC
[IBSC0004823], image seen).
Distribution. China: Sichuan (Muli Xian), Yunnan (Yongsheng Xian).
Oreocharis billburttii Mich.Moller & W.H.Chen, nom. nov. — Briggsia aurantiaca
B.L.Burtt, Notes Roy. Bot. Gard. Edinburgh 21: 237 (1955). TYPE: China, Xizang,
Anew Gesneriaceae genus and the transfer oiBriggsia species
201
Kongbo, Molo, Lilung Chu, 3050 m, 26 Jun 1938, Ludlow, G. Sherriff & G. Taylor
5670 (holotype BM [BM000041735], image seen).
Distribution. China: S Gansu, N Sichuan, W Yunnan, Xizang; N Myamnar.
Etymology^ To commemorate B.L. Burtt’s contribution to the taxonomy of the genus
Oreocharis.
Oreocharis elegantissima (H.Lev. & Vaniot) Mich.Moller & W.H.Chen, comb. nov.
— Didissandra elegantissima H.Lev. & Vaniot, Compt. Rend. Assoc. Fran^. Avancem.
Sci. 34: 425 ( 1 906). — Didymocarpus elegantissimus (H.Lev. & Vaniot) H.Lev., Compt.
Rend. Assoc. Fran?. 34: 428 (1906), nom. inval. — Briggsia elegantissima (H.Lev. &
Vaniot) Craib, Notes Roy. Bot. Gard. Edinburgh 1 1 : 265 (1919). TYPE: China, Kouy-
Tcheou [Guizhou], Pin-Fa, J. Cavalerie 239[h] (lectotype E! [E00387550], designated
here; PE [PE00030663] n.v.].
Distribution. China: Guizhou (Pingfa Xian, Dushan Xian).
Oreocharis latisepala (Chun ex K.Y.Pan) Mich.Moller & W.H.Chen, comb. nov.
— Briggsia latisepala Chun ex K.Y.Pan, Acta Phytotax. Sin. 26: 454 (1988). TYPE:
China, Zhejiang, Yunhe, 10 Oct 1932, S. Chen 837 (holotype IBSC [IBSC0649550],
image seen).
Distribution. China: Zhejiang (Yunhe Xian).
Oreocharis parva Mich.Moller & W.H.Chen, nom. nov. — Briggsia humilis K.Y.Pan,
Acta Phytotax. Sin. 26: 453 (1988). TYPE: China, Hubei, Lichuan, alt. 1300 m, 27
Aug 1975. G.R. Huang 3535 (holotype HMDB, n.v.; isotypes PE [PE00030685, image
seen], HIB [HIB0087078] n.v.).
Distribution. China: W Hubei.
Etymology. Named to reflect its small size.
Oreocharis parvifolia (K.Y.Pan) Mich.Moller & W.H.Chen, comb. nov. — Briggsia
paiwifolia K.Y.Pan, Acta Phytotax. Sin. 26: 457 (1988). TYPE: China, Guizhou, sine
loco,/. Cavalerie 3122 (holotype K! [K00085 8099]; isotype E! [E00135151]).
Distribution. China: Guizhou.
202
Gard. Bull. Singapore 66(2) 2014
Oreocharis pinfaensis (H.Lev.) Mich.Moller & W.H.Chen, comb. nov. — Didissandra
pinfaensis H.Lev., Repert. Spec. Nov. Regni Veg. 9: 328 (1911 ). — Briggsia pinfaensis
(H.Lev.) Craib, Notes Roy. Bot. Gard. Edinburgh 11: 264 (1919). TYPE: China, Kouy-
Tcheou [Guizhou], Pin-Fa, moist rocks, J. Cavalerie 3315 (lectotype E! [E00265025],
designated here; isolectotype K! [K000858091]).
Distribution. China: Guizhou (Pingfa Xian).
Oreocharis shweliensis Mich.Moller & W.H.Chen, nom. nov. — Briggsia forrestii
Craib, Notes Roy. Bot. Gard. Edinburgh 11: 237 (1919). TYPE: China, Yunnan,
Shweli-Salwin divide, alt. 8-9,000 ft., fl. & fr. Jun 1918, G. Forrest 17,552 (lectotype
E! [E00096870], designated here; isolectotypes Kn.v., PE [PE00030661] n.v.).
Distribution. China: Yunnan (Ruili Xian).
Etymology. Named after a collection locality, the Shweli valley.
Oreocharis tongtchouanensis Mich.Moller & W.H.Chen, nom. nov. — Briggsia
mairei Craib, Notes Roy. Bot. Gard. Edinburgh 1 1: 239 (1919). TYPE: China, Yunnan,
Tong-tchouan [Dongchuan], alt. 2700 m, fl. Sept, E.E. Maire 213 (lectotype E!
[E00396440], designated here; IBSC [IBSC0004831], image seen).
Distribution. China: Yunnan (Dongchuan Xian).
Etymology. Named after the old spelling of the collection locality, Dongchuan.
ACKNOWLEDGEMENTS. We are grateful to John McNeill for advice on nomenclatural
issues; Robert Mill for checking the Latin names; and Larry Skog and Goro Kokubugata for
their comments on the manuscript. We greatly acknowledge support by the National Natural
Science Foundation of China (grant nos. 31000258 and 31470306), the Open Project of the
Key Laboratory for Plant Diversity and Biogeography of East Asia, Kumning Institute of
Botany (grant no. KLBB201304), and the Chinese Academy of Sciences Visiting Professorship
Scheme. The Royal Botanic Garden Edinburgh is supported by the Rural and Environment
Science and Analytical Services division (RESAS) in the Scottish Government.
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Appendix. List of Briggsia names with their current status (bold).
Briggsia acutiloba K.Y.Pan = Oreocharis acutiloba (K.Y.Pan) Mich.Moller & W.H.Chen
Briggsia agnesiae (Forrest ex W.W.Sm.) Craib = Oreocharis agnesiae (Forrest ex WWSm.)
Mich.Moller & W.H.Chen
Briggsia amabilis (Diels) Craib = Loxostigma kurzii (C.B. Clarke) B.L.Burtt
Briggsia aurantiaca B.L.Burtt = Oreocharis billburttii (B.L.Burtt) Mich.Moller & W.H.Chen
Briggsia beauverdiana (H.Lev.) Craib = Briggsiopsis delavayi (Franch.) K.Y.Pan
Briggsia cavaleriei (H.Lev. & Vaniot) Craib = Loxostigma cavaleriei (H.Lev. & Vaniot)
B.L.Burtt
Briggsia chienii Chun = Oreocharis chienii (Chun) Mich.Moller & A.Weber
Briggsia crenulata Hand.-Mazz. = Oreocharis rosthornii var. crenulata (Hand.-Mazz.) Mich.
Moller & A.Weber
Briggsia damingshanensis L.Wu & B.Pan = Loxostigma damingshanensis (L.Wu & B.Pan)
Mich.Moller & H. Atkins
Briggsia delavayi (Franch.) Chun = Briggsiopsis delavayi (Franch.) K.Y.Pan
Briggsia dongxingensis Chun ex K.Y.Pan = Loxostigma dongxingensis (Chun ex K.Y.Pan)
Mich.Moller &Y.M.Shui
Briggsia dulongensis H.Li, nom. inval.
Briggsia elegantissima (H.Lev. & Vaniot) Craib = Oreocharis elegantissima (H.Lev. & Vaniot)
Mich.Moller & W.H.Chen
Briggsia forrestii Craib = Oreocharis shweliensis (Craib) Mich.Moller & W.H.Chen
Briggsia f itschii (H.Lev. & Vaniot) Craib = Glabrella mihieri (Franch.) Mich.Moller &
Y.M.Shui
Briggsia hians Chun = Oreocharis rosthornii (Diels) Mich.Moller & A.Weber
Briggsia humilis K.Y.Pan = Oreocharis parva (K.Y.Pan) Mich.Moller & W.H.Chen
Briggsia kurzii (C.B. Clarke) W.E.Evans = Loxostigma kurzii (C.B. Clarke) B.L.Burtt
Briggsia latisepala Chun ex K.Y.Pan = Oreocharis latisepala (Chun ex K.Y.Pan) Mich.Moller
& W.H.Chen
Briggsia longicaulis W.T.Wang & K.Y.Pan = Loxostigma longicaule (W.T.Wang & K.Y.Pan)
Mich.Moller & Y.M.Shui
Briggsia longifolia Craib = Oreocharis longifolia (Craib) Mich.Moller & A.Weber
Briggsia longifolia var. multiflora S.Y.Chen ex K.Y.Pan = Oreocharis longifolia var. multiflora
(S.Y.Chen ex K.Y.Pan) Mich.Moller & A.Weber
Briggsia longipes (Hemsl. ex Oliv.) Craib = Glabrella longipes (Hemsl. ex Oliv.) Mich.Moller
& Y.M.Shui
Anew Gesneriaceae genus and the transfer oiBriggsia species
205
Briggsia mairei Craib = Oreocharis tongtchoiianemis (Craib) Mich.Moller & W.H.Chen
Briggsia mihieri (Franch.) Craib = Glabrella mihieri (Franch.) Mich.Moller & Y.M.Shui
Briggsia muscicola (Diels) Craib = Oreocharis muscicola (Diels) Mich.Moller & A. Weber
Briggsia parvifolia K.Y.Pan = Oreocharis parvifolia (K.Y.Pan) Mich.Moller & W.H.Chen
Briggsia penlopi C.E.C.Fisch. = Oreocharis muscicola (Diels) Mich.Moller & A. Weber
Briggsia pinfaensis (H.Lev.) Craib = Oreocharis pinfaensis (H.Lev.) Mich.Moller & W.H.Chen
Briggsia rosthornii (Diels) B.L.Buitt = Oreocharis rosthornii (Diels) Mich.Moller & A. Weber
Briggsia rosthornii var. crenulata (Hand.-Mazz.) K.Y.Pan = Oreocharis rosthornii var.
crenulata (Hand.-Mazz.) Mich.Moller & A. Weber
Briggsia rosthornii var. wenshanensis K.Y.Pan = Oreocharis rosthornii var. wenshanensis
(K.Y.Pan) Mich.Moller & A.Weber
Briggsia rosthornii var. xingrenensis K.Y.Pan = Oreocharis rosthornii var. xingrenensis
(K.Y.Pan) Mich.Moller & A.Weber
Briggsia speciosa (Hemsl.) Craib = Oreocharis speciosa (Hemsl.) Mich.Moller & W.H.Chen
Briggsia stewardii Chun = Oreocharis stewardii (Chun) Mich.Moller & A.Weber
Gardens’ Bulletin Singapore 66(2): 207-214. 2014
207
A new species and a new record of Boesenbergia
(Zingiberaceae) for Thailand
J.D. Mood^, J.R Veldkamp^ & L.M. Prince^
Ryon Arboretum, University of Hawaii, 3860 Manoa Road,
Honolulu, HI 96822, USA
zi ngiber@ warml ava. com
^Naturalis Biodiversity Center, National Herbarium of The Netherlands,
P.O. Box 9517, 2300 RA Leiden, The Netherlands
Rhe Field Museum, Department of Botany, 1400 S Lake Shore Dr.,
Chicago, IL 60605, USA
ABSTRACT. Boesenbergia purpureorubra Mood & L.M.Prince is described and illustrated.
Boesenbergia longipes (King & Prain ex Ridl.) Schltr. is noted as a new record for Thailand
based on C Maknoi T38 (PSU, QBG).
Keywords. Boesenbergia, Thailand, Zingiberaceae
Introduction
The genus Boesenbergia Kuntze currently includes c. 82 species which are distributed
from SW India eastward to the Philippine Islands and south to the Wallace Line.
Although the majority of species are found within forests, some have adapted to
unusual habitats to include limestone fonnations near the ocean [Boesenbergia
ochroleuca (Ridl.) Schltr,] and sandstone mountain tops at over 1000 m [S. alba
(K.Larsen & R.M.Sm.) Mood & L.M.Prince]. As a consequence, Boesenbergia species
have evolved into a variety of vegetative fonns to adapt to specific environments.
They can be evergreen as are all the wild species in Borneo, or deciduous as are the
majority in Thailand, Plants can range from c. 10 cm tall [Boesenbergia panmla (Wall,
ex Baker) Kuntze] to over 130 cm (B. maxwellii Mood & L.M.Prince), forai single
clumps [B. curtisii (Baker) Schltr.] or develop into large, spreading populations of
connected stems (B. kingii Mood & L.M.Prince). Florally, Boesenbergia has evolved
to accommodate pollinators within each habitat, resulting in several different labellum
forms, from flat to very saccate. These variations aside, most Boesenbergia are easy
to identify by their basipetalous flowering sequence — the first flower opens near the
inflorescence apex and each subsequent flower, closer to the base.
In Thailand there are over 26 species currently recorded (Ridley, 1 899; Sirimgsa,
1987, 1992; Larsen, 1993, 1997; Saensouk & Larsen, 2002; Kharukanant & Tohdam,
2003; Mood et al., 2013, 2014). During preparation for a revision of Boesenbergia
for the Flora of Thailand, the authors compared the available Thai specimens to the
protologues and types of the c. 26 species. Surprisingly, the Thai taxon considered as
208
Card. Bull. Singapore 66(2) 2014
Boesenbergia longipes (King & Prain ex Ridl.) Schltr. (Sirirugsa, 1992) did not match
the type, Wray 4220 (K), the protologue (Ridley, 1899) or the expanded description
of Holttum (1950). To confirm the description of Sirirugsa, c. 10 similar specimens
collected in Thailand and annotated as “-5. longipes'' were examined (AAU, BK, BKF,
QBG), It was evident that the vegetative fonn of the Thai taxon varied considerably
from Wray 4220. Additionally, flowering plants observed in Peninsular Thailand,
near locations as noted on the specimens, did not match Ridley or Holttum ’s floral
descriptions. Since this taxon showed no similarity to any other known Boesenbergia,
it is here considered a new species and is described as Boesenbergia purpureorubra.
During examination of specimens, Maknoi T38 (PSU, QBG), an unidentified
Boesenbergia from Yala Province on the Malaysian border, was found to match the
type and protologue of B. longipes (King & Prain ex Ridl.) Schltr., thus making it a
new record for Thailand.
Taxonomy
Boesenbergia purpureorubra Mood & L.M.Prince, sp. nov.
Boesenbergiae pulcherrimae (Wall.) Kuntze similis, planta minore c. 40 cm alta, caule
breviore c. 6 cm longo pagina corrugatissima atro purpureorubro, petiolis longioribus
c. 3-8 cm longis differt. TYPE: Cultivated in Hawaii, USA, 1 August 2014, Mood
14P19 (holotype BKF; isotype AAU). Originally from Thailand, Ranong Province,
E. of Khao Niwet, 09° 57.506'N 98°39.134'E, 880 m asl, evergreen forest, 24 August
2011, Mood & Vatcharakorn 3106, cultivated as M3 106 (Fig. 1-3)
Boesenbergia longipes auct. non Schltr.: Sirirugsa, Nat. Hist. Bull. Siam Soc. 40: 76
(1992).
Deciduous^ perennial herb up to c. 40 cm tall, forming tight clumps. Rhizome with
multiple elements developing from the base of the previous element, c. 5 cm long, c.
8 mm diam. in a mature plant with five stems, internally and externally white (young),
internally dark puiple (mature); roots many, fleshy, to c. 20 cm long, 3-4 mm diam.,
externally and internally white or pink, covered in short root hairs along the full length,
apex expanded, fusifomi, c. 4-5 cm long, 5-10 mm diam., surface smooth, root hairs
along the full length, few fibrous roots. Stems numerous, increasing in number each
season, to 6 cm long, c. 7 mm diam., light green, glabrous, 1 or 2 leafless sheaths,
5-1 1 X 2.5 cm, deeply corrugate, solid or spotted purple-red, glabrous. Leaves 2 or
3, upper two nearly opposite; leaf sheaths c. 11-14 cm long, light green, sparsely
sericeous, corrugate, covered in part by the leafless sheaths; ligule bilobed, up to 17
mm, lobes oblong to lanceolate, green, few hairs; petiole 3-8 cm long, 2 mm wide,
light green or with purple-red spots, sparsely sericeous; lamina elliptic, sometimes
asymmeti'ic, 18 x 8 cm (lower) to 25 x 8 cm (upper), base attenuate to rounded, often
oblique, apex acuminate, adaxially dark, glossy green, veins prominent, few long hairs.
Anew species and record oi Boesenbergia for Thailand
209
Fig. 1. Boesenbergia purpureorubra Mood & L.M.Prince. Ink line drawing with watercolour
of the type plant. (Drawn by Linda Ann Vorobik).
210
Card. Bull Singapore 66(2) 2014
Fig. 2. Boesenbergia purpureorubra Mood & L.M.Prince. A. Mature plant in flower. B.
Flowers in side and semi-side view. C. Flower in front view. D. Inflorescence (scale in cm).
(Photos: J. Mood of M3 106)
Anew species and record oi Boesenbergia for Thailand
211
Fig. 3. Boesenbergia purpureorubra Mood & L.M.Prince. A. Flower dissection. B.
Inflorescence with bracts spread showing their attachment. C. Anther and stigma (mature). D.
Underground architecture (rhizome, roots, tuberous roots). (Photos: J. Mood of M3 106)
212
Card. Bull. Singapore 66(2) 2014
abaxially purple, dark red or green, sericeous. Inflorescence terminal, basal y4 clasped
between the leaf sheaths, apical % exposed; peduncle c. 1 cm long, 4 mm diam., white,
glabrous; spike fusifoi*m, c. 9-13 cm long, c. 1 cm wide at the middle, c. 7 mm thick,
longitudinally furrowed on the flowering side; bracts 16-24, narrowly ovate, c. 35
X 6 mm, light green, sericeous, compressed together radially on the flowering side,
the rachis exposed on the non-flowering side; bracteole nan'owly ovate, open to the
base, c. 32 X 10 mm, white, glabrous, on the opposite side of the floral tube from the
bract. Flowers c. 3.5 cm long, labellum oriented toward the bract apex, all deflexed
downward; calyx tubular, c. 7 mm long, light green with red dots, few hairs, apex
undulate, with 2-3 lobes; /?ora/ tube 30-35 mm long, 1 .5 mm diam. at the base, white,
glabrous; dorsal corolla lobe c. 1 2 x 5 mm, slightly convex, oblong, white, glabrous,
apex cucullate, lateral corolla lobes c. 15 x 6 nun, oblong, white, glabrous, apex
cucullate; androecial cup c. 4 mm long, c. 4 mm wide, oriented c. 90” to the floral
tube, throat mostly glabrous; labellum deeply saccate, elongate to oval (natural shape),
c. 2.8 X 1 .6 cm, throat glossy dark red with transverse red bars on the sides and red
spots in the centi'e, apical area glossy violet-pink, lighter and slightly transparent on
the margin, internally glabrous, externally with short, glandular hairs, apex rounded,
slightly undulate, deflexed; lateral staminodes obovate, c. 10 x 8 mm, yellowish-
white, externally covered in short, glandular hairs, apex tmncate to rounded. Stamen
c. 10 mm long, filament c. 2 x 1 mm, white; anther c. 8 x 3 nun, connective tissue with
glandular hairs from the base up Ya the length, apical part glabrous, anther thecae c. 7 x
1.5 mm (each), dehiscent tluoughout the full length, white, anther crest absent, pollen
white. Ovfl/jtrilocular, cylindric, c. 3 x 2.5 mm, light green, glabrous; style filiform, c.
3.7 cm long, white; stigma orbicular, white, ostiole circular, ciliate; epigynous glands
linear, two, c. 3 lum long, white. Fruit not seem (Measurements based on living,
cultivated material of M3 106).
Distribution. Peninsular Thailand.
Ecology. This species is found in evergreen forest between 50-300 m on soils derived
from granite or sandstone.
Phenology’. In cultivation in Hawaii, flowering begins in late June and continues
through October. Flowers open in the morning and close early the following day.
Observations in the wild are similar.
Etymology. Named for the purple-red colour of the leafless sheaths and lamina
underside.
Additional specimens examined: THAILAND. Ranong: Kapis, Kampon, 100 m, 20 Nov 1973,
T. Sandsuk 633 (BKF); 10 km south of Khao Khai, 2 Aug 2013, J. Mood & P Vatcharakorn
3376 (AAU); Kampuan, 11 Jul 1999, C. Maknoi 53 (QBG). Surat Thani: Ban Tung Tao, 4
A new species and record of Boesenbergia for Thailand
213
Aug 1927, Klein? 13153 (BK); Khao Sok, 25 Oct 1990, K. Larsen 40887, Phang Nga: 8 km
north of Bang Wan, 2 Aug 2013, J. Mood & P. Vatcharakorn 3380 (AAU); Bangwan Stream,
14 Aug 2006, T. Muadsub 116 (PSU, BKF). Trang: Khao Chong, 200 m., 12 Aug 1975, J.F.
Maxwell 75-797 (BK); Muang Dist., Khao Chong N.P., 14 Jul 1985, P. Sirirugsa 1021 (PSU,
BKF); Khao Chong forest, 150 m, 30 Oct 1984, J.F. Maxwell 84-375 (BKF). Narathiwat:
Waeng, 50 m, 23 Aug 2006, M. Poopath 23 (BKF).
Notes. It is not known how this ginger initially came to be identified as Boesenbergia
longipes, as the species was originally described from Perak, Malaysia (Ridley, 1 899).
The type, Wray 4220 and a similar specimen Corner S.F.N. 31673, cited by Holttum
(1950), are quite distinct from the Thai taxon. Some of the vegetative differences for
the Perak plants are the loose, spreading leaf sheaths with short stem, long petioles and
inflorescence close to the ground. Boesenbergia purpureorubra has short leaf sheaths
tightly clasped, short petioles and an inflorescence at the top of the plant. Ridley
named Wray’s plant “long foot”, noting the “rhizome rather far creeping”. The new
species is a very tightly clumped plant with multiple stems from a short rhizome. A
noticeable character of the new taxon is its deeply corrugated, dark purple-red leafless
sheaths. The underside of the lamina is also the same colour even when mature. These
characters are quite noticeable on the Thai specimens such as Maxwell 75-797 (BKF).
Ridley described the flower of Boesenbergia longipes has having an oblong,
crisped and thickened labellum, a rounded apex and a thick, central bar. Colour notes
were not included. Flolttum (1950) broadened the description and provided colour
notes based on a flower in alcohol from Corner S.F.N. 31673, collected near the type
locality. Flolttum mentions that since the Comer flower was larger than the type (which
was only a broken portion) he was not sure if “one or two species exist.” It is the
experience of the first author that flower size can be quite variable based on the vigour,
age of the plant and population variation. More important is that both Wray’s type and
Corner’s specimen closely match in overall morphology.
Recently, by request of the authors, the type locality was visited by Lim Chong
Keat (Penang, Malaysia) in order to ascertain whether Boesenbergia longipes was
still extant in this area. The species was found and the plants matched the vegetative
morphology of the descriptions and specimens, while the floral stmcture and colours
were exactly as Flolttum described. With this added validation it is quite clear that
Boesenbergia longipes is strikingly different from the Thai taxon described here as B.
purpureorubra (Table 1).
True Boesenbergia longipes has, however, once been collected in Thailand as
recently discovered by the first author. The specimen, C. Maknoi T38 (QBG, PSU),
annotated as Boesenbergia sp., was collected near the Malaysian border in Yala
Province, Betong, Ban Chulapom Pattana 10, 5 August 1999. The forest is evergreen,
similar to the type locality in Perak. The collector noted, “Lip white with yellow
median band and red lines on either side from base to half of length” matching Floltum
(1950). This specimen constitutes a new record for Thailand.
214
Card. Bull. Singapore 66(2) 2014
Table 1. Comparison of Boesenbergia purpureorubra and 5. longipes
Character
B. purpureorubra (M3 106)
B. longipes (M3 106)
Plant height
40 cm
c. 60 cm
Petiole length
3-8 cm
to 23 cm
Lamina
18-25 X 8 cm
30 X 10 cm
Inflorescence
9-13 cm long
5 cm long
Floral tube
3-3.5 cm
4.5 cm
Labellum
2.8 cm long, oval,
2 cm long, obovate.
saccate
non saccate
Labellum colour
white, red, violet-pink
white, red, yellow median band
Filament
2 mm
4 mm
Anther
7 mm
4 mm
Anther crest
none
oblong, rounded, fleshy
ACKNOWLEDGEMENTS. We thank the staff at BK, BKF and QBG for assistance; The
Field Museum, Chicago, USA (F) for use of laboratoty facilities; Lim Chong Keat (Penang,
Malaysia) for photography; and Linda Ann Vorobik (Berkeley, USA) for the watercolour.
References
Holttum, R.E. (1950). The Zingiberaceae of the Malay Peninsula. Gard. Bull. Singapore 13(1):
1-249.
Kharukanant, B. & Tohdam, S. (2003). Anew ^^Qcio^of Boesenbergia O. Kuntze (Zingiberaceae)
from Peninsular Thailand. Folia Malaysiana 4: 19-24.
Kuntze, O. (1891). Scitaminaceae. Revisio Generum Plantarum 2; 682-698. Leipzig: Felix.
Larsen, K. (1993). Boesenbergia tenuispicata (Zingiberaceae): a new species from Thailand.
Nord, J. Bot. 13: 281-283.
Larsen, K. (1997). Further studies in the genus Boesenbergia. Nord. J. Bot. 17: 361-366.
Mood, J.D., Prince, L.M., Veldkamp, J.F. & Dey, S. (2013). The history and identity of
Boesenbergia longijiora (Zingiberaceae) and descriptions of five related new taxa.
Gard. Bull. Singapore 65: 47-95.
Mood, J.D., Veldkamp, J.F., Dey, S. & Prince, L.M. (2014). Nomenclatural changes in
Zingiberaceae: Caulokaempferia is a superfluous name for Monolophus mdJirawongsea
is reduced to Boesenbergia. Gard. Bull. Singapore 66(2): 215-231.
Ridley, H.N. (1899). The Scitamineae of the Malay Peninsula, J. Straits Branch Roy. Asiat.
Soc. 43: 85-184.
Saensouk, S. & Larsen, K. (2002). Boesenbergia baimaii, a new species of Zingiberaceae from
Thailand. Nord. J. Bot. 21: 595-597.
Sirirugsa, P. (1987). Three new species and one new combination in Boesenbergia
(Zingiberaceae) from Thailand. Nord. J. Bot. 7: 421-425.
Sirirugsa, P. (1992). A revision of the genus Boesenbergia Kuntze (Zingiberaceae) in Thailand.
Nat. Hist. Bull. Siam Soc. 40: 67-90.
Gardens’ Bulletin Singapore 66(2): 215-231. 2014
215
Nomenclatural changes in Zingiber aceae:
Caulokaempferia is a superfluous name for Monolophus
and Jirawongsea is reduced to Boesenbergia
J.D. Mood^, J.R Veldkamp^, S. Dey^ & L.M. Prince^
'Lyon Arboretum, University of Hawaii, 3860 Manoa Road,
Honolulu, HI 96822, USA
zmgiber@ wamilava. com
^Naturalis Biodiversity Center, National Herbarium of The Netherlands,
P.O. Box 9517, 2300 RA Leiden, The Netherlands
-^Department of Botany, Nagaland University, Lumami 798627,
Zunheboto, Nagaland, India
Rhe Field Museum, Department of Botany, 1400 S Lake Shore Dr.,
Chicago, IL 60605, USA
ABSTRACT. Wallich published Monolophus Wall. (Zingiberaceae) in 1832 for two taxa he
had described previously in 1820 as Kaeinpferia linearis Wall, and K. secunda Wall. He also
mentioned M. ? elegans Wall., but the “?” indicates that he was not certain that this species
belonged to this new genus. Consequently, elegans., while validly published, cannot
be the lectot^pe of the generic name. In 1 964 Larsen argued against accepting Monolophus and
established the alternative Caulokaempferia K.Larsen, Caulokaempferia, typified by the name
C. linearis (Wall.) K.Larsen, is, however, supeiRuous. Monolophus is hereby reinstated with
22 new combinations. Its phylogenetic position is shown in relation to other genera. Based
on comparative nuclear and chloroplast DNA sequence data analyses, Jirawongsea Picheans.
(previously Caulokaempferia, pro parte) is reduced to Boesenbergia Kuntze with five new
combinations.
Keywords. Kaempferia, molecular phylogeny, nomenclature, taxonomy
Introduction
Over the past several years the authors have been collecting molecular data from both
the nuclear ITS and the chloroplast trnK intron to produce a phylogeny of the genus
Boesenbergia Kuntze. In the process, other Zingiberaceae genera have been included as
outgroups. Caulokaempferia K.Larsen is one of these genera. Until recently, this genus
consisted of 29 species described from the Lao P.D.R., Northeast India and Thailand.
In 2007, a molecular phylogeny of Caulokaempferia using samples of 23 taxa from
Thailand and the Lao P.D.R. was completed (Chaiyoot, 2007). The results showed the
genus to be polyphyletic with two distantly related clades. Since a sample of the type
species, Caulokaempferia linearis, was not included, no definitive conclusion was
216
Card. Bull. Singapore 66(2) 2014
possible as to which clade should retain the name Caulokaempferia. Picheansoonthon
et al. (2008) used these results in combination with the morphological characters of
flower colour and capsule type (Larsen, 2003) to detemiine that the largest clade (18
species) represented tme Caulokaempferia, while the smaller clade (five species) was
better treated as a new genus, which was called Jirawongsea Picheans., despite the
samples clustering within the Boesenbergia clade. Also, a morphological comparison
of Jirawongsea to Boesenbergia was not included.
When conducting a brief historical review of Caulokaempferia as part of this
study it was discovered that an earlier name, Monolophus Wallich (1832) appeared
to have been validly published with two species, one of which is the type of
Caulokaempferia. This intriguing infonnation triggered an in-depth nomenclatural
study to understand why Monolophus had not been used in lieu of Caulokaempferia.
This paper provides new insights into the nomenclature, phytogeny and
morphology of Caulokaempferia and Jirawongsea.
While all four authors made significant contributions to the manuscript, JDM
is the key investigator and general manager of this paper, JFV provided the historical
research as a basis for nomenclatural decisions, SD was instrumental in making field
observations, and LMP provided supporting molecular phylogenetic research.
Nomenclature
Wallich (1820) described two new species of Kaempferia L. (Zingiberaceae): K.
linearis Wall, and K. secunda Wall. At the time he thought them to be atypical for
this genus and gave a brief diagnosis for this “group”, but not a rank, nor a name:
“Caulescent, with an entire cresf ’.
In 1829 he added Kaempferia elegans Wall, and stated, “[it] belongs to the
section which 1 have long ago [1820] indicated ... All three ought perhaps to be
removed from Kaempferia, and fonned into a distinct genus for which 1 would propose
the name Monolophus [single, entire crest].” Monolophus is here a provisional name
for both the section and the genus and is, therefore, invalid (McNeill et al., 2012 - Art.
36.1b) although there is, yet again, a diagnosis: “caulescent habit, absence of tubers
and entire crest”.
The nomenclatural articles cited here are according to the International Code of
Nomenclature (TCN) (McNeill et al. 2012).
Wallich (1 832) p\xh\is\{Q& Monolophus and included three species with references
to earlier publications. Although Monolophus as a genus is not separately mentioned.
Art. 35.1 and Ex. 1 on Suaeda Forssk. do not apply because the name of the genus and
its three species are validly published through direct reference to previous diagnoses
and descriptions (Art. 38.1). Monolophus linearis andM secunda are clearly syntypes
while M. elegans is not, as it was cited with a query and was not part of the original
concept in 1820.
Nomenclatural changes in Zingiberaceae
217
As Wallich (1829) so beautifully illustrated Kaempferia elegans it became the
best known species of the assemblage, thus causing later problems in lectotypification
(e.g. supposedly by Endlicher (1837)).
Monoloplms was recognised by Endlicher (1837), followed by Steudel
(1841), Horaninov (1862), Pfeiffer (1874), and more recently by Wu & Chen (1978).
Endlicher ’s citation of Kaempferia elegans is not to be regarded as a lectotypification,
but as a representative of the genus. As Kaempferia elegans is not a syntype this cannot
be a lectotype, yet it was eiToneously accepted by Larsen (1964: 165), followed by
Buitt & Smith (1972: 216), Wu & Chen (1981), and Newman et al. (2004: 118), and
thus apparently by the Index Nominum Genericorum (Farr, 2013).
Horaninov (1862) is sometimes cited as the validating author of Monolophus
but from the text it is clear that he was not, as he refers to Endlicher (1837), Roxburgh
(1820) and Wallich (1832).
Bentham (1883) regarded Monolophus as connecting Kaempferia L. sections
Stachyanthesis Benth. and Soncorus Rumph. ex Horan. The latter is an invalid name for
Kaempferia sect. Kaempferia as an autonym is required. He included only Kaempferia
linearis and K secunda and did not mention K elegans.
Baker (1892) accepted Monolophus as a subgenus of Kaempferia and included
seven species, some of which are now placed elsewhere, e.g. in Boesenbergia Kuntze,
Camptandra Ridl., Kaempferia and Stahlianthus Kuntze. He was more or less followed
by Schumann (1904: 73) who created a later isonym, as he eiToneously thought that
Baker had regarded it as a section.
Larsen (1964) transfQrrQd Kaempferia linearis andK. secunda to a new genus,
Caulokaempferia with C. linearis as the type. This transfer was supported by then new
morphological findings and cytological data.
As was argued above, Larsen (1964) was incorrect in regarding Kaempferia
elegans as the type of Monolophus, which he saw as more related to K galanga L., the
lectotype of Kaempferia. He, therefore, XQ]QciQd Monolophus because 1 ) the type species
would be a Kaempferia and 2) “the name refers to a character widely distributed also
in neighbouring genera”. The first argument is to be rejected as Kaempferia elegans is
not a candidate for lectotypification. Tlie second is a taxonomic, not a nomenclatural,
argument and also contravenes Art. 51.1 (a name is not to be rejected because it would
be inappropriate).
Because Kaempferia linearis and K. secunda are the syntypes of Monolophus,
Larsen (1964) created a supeiTluous name. The fact that, in 1964, Monolophus had not
yet been lectotypified is irrelevant (Art. 52.2a). Monolophus linearis Wall, was later
designated as the lectotype by Wu & Chen (1978: 28).
Even though Wu & Chen (1978) recognised the validity of Monolophus they
later (Wu & Chen, 1981) changed their opinion on the validity of Monolophus and
reverted to the name Caulokaempferia which has been in general usage to the present
day.
Here we reinstate Monolophus as the valid name for this genus and provide 22
new combinations (Appendix 1).
218
Card. Bull. Singapore 66(2) 2014
Materials and Methods
Molecular phylogeny. Caulokaempferia and Jirawongsea leaf tissue samples were
obtained from living plants (where available) and dried in silica gel. Supplemental
ingroup data were obtained from GenBank, as were additional sequences which
provided the scaffold for the taxa sampled here. A complete list of samples and
GenBank accession numbers are provided in Appendix 4. The Caulokaempferia
nucleotide data produced by Chaiyoot (2007) have not yet been deposited into any
of the four international sequence repositories BOLD, DDBJ, EMBL, GenBank, thus
they could not be included here.
DNA extraction follows Kress et al. (2002) while amplifr cation and analytical
methods follow Mood et al. (2013). Only minimal detail is provided here. The nuclear
ribosomal ITS (nrlTS) region was amplifred using the 18S-F and 26S-R primers
(Prince, 2010), The plastid trnK region was amplifred in two parts, the first using IF
and 1235R primers and the second using mlF and 2R. Data were collected on an
ABI Genetic Analyzer and sequences of each specimen were edited and a consensus
sequence was generated in Sequencher v4.9 (Gene Codes Corporation, Ann Arbor,
Michigan, USA). Sequences were aligned manually in Se-al (Rambaut, 1996) and
areas of ambiguous aligmnent identified. Data were analysed under parsimony criteria
by genomic data partition, frrstly independently and later in combination. Data were
also analysed including and excluding ambiguously aligned regions to determine if this
altered the resulting tree topologies. Maximum parsimony analyses were conducted in
PAUP* (version 4.0b 10; Swofford, 2002). Heuristic search methods were conducted in
each case with 1000 random addition replicates. Branch support was estimated using
parsimony bootstrap (BS) in PAUP*.
Results
Both the individual and combined data analyses agreed in topology. All trees from
all analyses agreed (or did not conflict) with a monophyletic Caulokaempferia
(Monolophus) and Boesenbergia. Additionally, both phytogenies placed all samples of
Jirawongsea within Boesenbergia, For brevity, only the results of the combined data
analyses are shown and discussed below. The combined ITS and trnK data analysis
(ambiguously aligned regions excluded) produced 64 shortest parsimony trees, one
of which is shown in Figure 1. All samples of Jirawongsea (labelled Boesenbergia
in Fig. 1) are part of a strongly supported (BS=100%) clade that includes B. rotunda
(L.) Mansf and B. curtisii (Baker) Schltr., which is nested within a monophyletic
Boesenbergia (BS=56%). These findings are consistent, whether ambiguously aligned
data are included or excluded from the analyses. Analyses of just the trnK or the ITS data
partitions recover similar tree topologies. The only differences in topology generally
involve minor ehanges in sister taxon relationships (with low support) among genera
of Zingibereae. Branch support is generally lower in the separate analyses.
Nomenclatural changes in Zingiberaceae
219
51
98
93
Boesenbei-gia alba M3253*
Boesenbergia alba M 1 2C30*
Boesenbergia violacea *
Boesenbergia thailandica
Boesenbergia burttii*
Boesenbergia rouinaa
Boesenbergia cnnisii*
Boesenbergia plieata lurida
Boesenbe rgia pulcherrirna'^
Boesenbergia hamUtonii
Comukaempferia aurantiflora
Zingiber graniineum
DisticlwchJamys rub ro striata
Kaempferia rotunda
Haniffia albiflora
Pomtnereschea lackneri
Rbynchanthiis beesianus
Roscoea purpurea
Hitchenia glauca
Ryrgophyllum ytinnanense
Camptandra parvida
Hedychiuin botdeloniannm
Ilemtorchis rhodorrhachis
100
100
Monolopluis coenobialis
Monoloplius saxicola *
Monulopinis limianus*
Monoloplius linearis
Monoloplius sikkiinensis*
Globha curtisii
Hornsiedtia hainanensis
Siliquamomum tonkinense
Tamijia flagellaris
Siphonocitilus kirkii
10 changes
Fig. 1. Phylogram of 1 of the 64 shortest maximum parsmiony trees for Boesenbergia and
Monoloplius (Zingiberaceae) based on an analysis of combined ITS and trnK sequence data.
^ indicates the type species for the genus. * indicates sequences generated for this study.
Numbers above branches are bootstrap support values. Bold branches were recovered in the
strict consensus tree.
Discussion
Larsen (1964) described Caulokaempferia with eight species that he considered to be
yellow-flowered. Larsen & Smith ( 1 972) added Caulokaempferia alba (white-flowered)
and the yellow-flowered, C. yunnanensis (Gagnep.) R.M.Sm. (See Appendix 2). The
inclusion of these taxa was problematic, as they were quite different from each other
and the type. The authors discussed the diversity of Caulokaempferia and Boesenbei^ia
and suggested, on balance, that these species belonged in Caulokaempferia but
conceded that characters such as the “large concave bracts ... [with a] lamina-like
extension of the apex.” in C. yunnanensis were more complex and required extending
the generic limits. Recent sampling of this taxon in molecular studies by Kress et al.
(2002), Zaveska et al. (2012) and here (Fig. 1), has shown it to have close affinity
to Curcuma L., but we maintain it in Pyrgophyllum (Gagnep.) T.L.Wu & Z.Y.Chen.
Larsen & Smith (1972) considered the flower of Caulokaempferia alba to be similar
in appearance to other species in Caulokaempferia, but its vegetative form, trilocular
capsule and flowering sequence required considerable accommodation. In this species
the flowers open sequentially, top to bottom, whereas in the type species, they open
bottom to top. Obviously, this was a disconcerting fact, as Larsen & Smith ( 1 972) stated
220
Card. Bull. Singapore 66(2) 2014
“Such a mode of flowering has not been previously observed m Caulokaempferia, and
the possibility of affinity to at least some Boesenbergia cannot be discarded.”
Based on the first author’s study of Boesenbergia across their range, it
appears that all of the species presently described can be distinguished from all other
Zingibereae (except Haplochorema K.Schum.) by their basipetalous flowering. This
is defined here as a flowering sequence where the first flower to open is at or near the
apex and subsequent flowers open sequentially down the rachis from single-flowered
bracts. In contrast, Caulokaempferia species are acropetalous with the flower-opening
sequence irom bottom to top. It should be noted that some Caulokaempferia have
circinni of 2^ flowers per bract, and in this case, the flowering sequence in the circinni
in relation to the overall sequence is undocumented.
Larsen & Smith (1972) mentioned several characters in Caulokaempferia alba
which they thought were atypical of Boesenbergia — a non-saccate labellum, a crested
anther and no [androecial] tube at the base of the labellum. In fact, these three characters
have been documented in Boesenbergia descriptions (Valeton, 1918; Holttum, 1950).
For example, Boesenbergia rotunda (Fig. 3C), a well-known ginger of ethnobotanical
importance, and B. curtisii (Fig. 3D) have these “atypical” characters. Perhaps Larsen
& Smith were referring to Boesenbergia species similar to the type, B. pulcherrima
(Wall.) Kuntze, which have a saccate labellum, an uncrested anther and an androecial
tube (Holttum, 1950).
Wallich (1820) did not note flower colour in the original description of
Kaempferia linearis and nor is a colour shown in the drawing received by Kew (K) on
October 1 7, 1 828 (annotated as “Wallich 1 828 East Ind. Co. #25”). The first mention of
the flower colour was by Baker (1892) who noted “flowers white ... tinged with yellow
at the tliroat”, but it is unknown where this information origmated. Contrary to Baker’s
colour description, Parryi 281 (K) from Assam ( 1 927) was annotated “flowers yellow”.
Larsen (1964) cited this specimen as representative of Caulokaempferia linearis, but
in its description he stated “ [flower] tinged with yellow at the throat”.
By 2003, five more Caulokaempferia species had been published by Larsen
(1973,2003). Two of these, Caulokaempferia thailandica and C. had characters
very similar to C. alba. It became apparent to Larsen that the accommodation of these
latter taxa [and Caulokaempferia alba] within Caulokaempferia was becoming more
problematic. To preclude moving these species to another genus, he taxonomically
grouped them into either yellow-flowered with unilocular capsules (to include the
type) or into “the whitish- to violet-flowered species” with trilocular capsules. He
further stated, “Until the relationships between these groups have been satisfactorily
elucidated thi*ough molecular studies, [1 have] chosen to keep the two groups united in
the genus Caulokaempferia'' (Larsen, 2003).
The first phylogenetic study to sample Caulokaempferia was by Kress et al.
(2002). Only three species from the yellow-flowered group were included. The results
showed affinity to Hanijfia albiflora K.Larsen & Mood. Ngamriabsakul et al. (2004)
and Thong-a-ram et al. (2005) sampled Caulokaempferia violacea, showing it to be
closely allied to Boesenbergia. Chaiyoot (2007) sampled both Caulokaempferia colour
groups, but without the type, C. linearis. His results showed the genus to be comprised of
Nomenclatural changes in Zingiberaceae
221
Fig. 3. A. Monolophus petelotii (K.Larsen) Veldk. & Mood. B. Boesenbergia alba (K.Larsen
& R.M. Smith) Mood & L.M.Prince. C. Boesenbergia rotunda (L.) Mansf. D. Boesenbergia
curtisii (Baker) Schltr. (Photos: A, J. Leong-Skomickova; B-D, J. Mood)
two unrelated clades. The five taxa of Larsen’s white-fiowered group clustered within a
strongly supported monophyletic Boesenbergia (BS=100%). This finding brought full
circle the observation of Larsen & Smith (1972) concerning the flowering sequence of
Caulokaempferia alba being the same as in Boesenbergia. Picheansoonthon et al. (2008)
proposed that these five taxa represented a new genus and named them Jirawongsea,
despite the findings of the molecular data analyses. Since the type, Caulokaempferia
222
Card. Bull. Singapore 66(2) 2014
linearis had not been included by Chaiyoot (2007), no direct detennination could
be made as to which clade actually represented Caulokaempferia. In lieu of this,
Picheansoonthon et al. (2008) followed the floral colour grouping of Larsen (2003) and
selected the yellow-flowered group as representative of Caulokaempferia. To support
their generic proposal, the taxonomic discussion was confined to the morphological
dissimilarities between Caulokaempferia and Jirawongsea with no mention of the
morphological similarities or phylogenetic affinity of the latter to Boesenbergia.
Figure 1 shows that Caulokaempferia linearis (as Monolophtis linearis) does
indeed group with the other species of Monolophus which are predominantly yellow-
flowered. Surprisingly, as can be seen from a photograph of Monolophus linearis
taken near the type locality in Meghalaya, India, the flower is not yellow, but white
(Fig. 2B) as was stated by Baker (1892). In fact, none of the Indian species are yellow-
flowered: Caulokaempferia secunda flowers are dark pink to violet (Fig. 2D) and
those of C. sikkimensis are white. Consequently, taxonomic grouping by flower colour
was an imperfect fit.
Based on phylogenetic and taxonomic investigations, Jirawongsea is reduced
to Boesenbergia with five new combinations (Appendix 3).
Since the most recent circumscription of Caulokaempferia (Larsen & Smith,
1972) included characters of taxa that are no longer in this genus, an updated
description of the genus is provided here under the name Monolophus.
Monolophus Wall., PI. Asiat. Rar. 1 : 24, t. 27 (1829), nom. prov. inval.; Wall., Numer.
List 6591-6593 (1832), validation; Wall, ex Endl., Gen. PI, 225 (1837), isonym. -
Kaempferia L. subg. Monolophus (Wall.) Wall, ex Baker in Hook.f , FI. Brit. India
6: 222 (1894); Baker ex K.Schum. in Engl., Pflanzenr. IV, 46, Heft 20: 73 (1904),
isonym. - Caulokaempferia K.Larsen, Bot. Tidsskr. 60: 166 (1964), nom. superfl.
TYPE: Monolophus linearis Wall., lectotype designated by Wu & Chen (1978).
Perennial herbs up to 45 cm; stems with 2-A bladeless sheaths at the base.
Leaves 3-10, sessile or petiolate, ligule small, bilobed to entire. Inflorescence
terminal, acropetalous, flowers opening singly (?). Bracts 1-10, distichous, lanceolate,
acuminate, the margins quite free to the base; \-A flowered. Bracteoles membranous,
often not associated with the first flower in those species with single-flowered bracts.
Calyx tubular, often 2-3 dentate, not deeply split unilaterally. Floral tube long, narrow,
widening at the mouth; lobes 3, the dorsal broader and a little longer than the laterals.
Lateral staminodes petaloid, normally small in relation to the labellum. Labellum
large, orbicular, entire or bilobed, slightly concave. Stamen usually with a very short
filament (occasionally up to 5 mm long) or sessile on the floral tube; anther thecae
mostly parallel, dehiscing longitudinally; anther crest conspicuous, entire or dentate,
often reflexed. Epigymous glands short, mostly linear, free Ifom each other. Ovary
unilocular; placentation free central. Fruit a unilocular capsule with a lateral suture.
Seeds numerous, small, ellipsoid, glabrous or with a dense indumentum, aril white.
Nomenclatural changes in Zingiberaceae
223
Fig. 2. A. Boesenbergia alba (K.Larsen & R.M. Smith) Mood & L.M.Prince. B. Monolophus
linearis (Wall.) Wall. C. Monolophus saxicola (K.Larsen) Veldk. & Mood. D. Monolophus
secundus (Wall.) Wall. (Photos: A & C, J. Mood; B & D, S. Dey)
ACKNOWLEDGEMENTS. We would like to thank Poonsak Vateharakom (Chanthaburi,
Thailand) for assistance in the field; R. Pooma (BKF) and L. Pashirajan, Botanical Survey
India (CAL) for botanical assistance; J. Leong-Skornickova (SING) for photography; PC.
Boyce (UNIMAS, Malaysia) for helpful comments; S.H. Chen (TAIE) for the translation of
Chinese texts; the reviewers for their time and diligence; and the Field Museum (F), Chicago,
USA, for use of laboratory facilities. Colour plates were created by T.D. Shafto (Hawaii, USA).
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Appendix 1. Enumeration of Monolophus species
1. Monolophus amplexicaulis (Suksathan) Veldk. & Mood, comb. nov. - Caulokaempferia
amplexicaulis Suksathan in K.Larsen et al., Nordic J. Bot. 23: 401, t. 1, 2. (2005)
tyamplexicaule’'). TYPE: Thailand, Mae Hong Son, Muang Distr., Doi Hua Kai-Doi Pui,
Wongnak et al. 705 (holotype QBG; isotypes AAU, BKF, E, K, US).
2. Monolophus appendiculatus (K.Laisen & Triboun) Veldk. & Mood, comb. nov. -
Caulokaempferia appendiculata K.Larsen & Triboun in K.Larsen, Nordic J. Bot. 22: 409, t. 1
(2003). TYPE: Thailand, Chiang Mai, Ang Kang, Kop, Dung village, Triboun 617 (holotype
AAU; isotypes BK, BKF).
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Card. Bull. Singapore 66(2) 2014
3. Monolophus bolavenensis (Picheans. & Koontemi) Veldk. & Mood, comb. nov. -
Caulokaempferia bolavenensis Picheans. & Koontenn, in Picheans. et al, Nat. Hist. Bull. Siam
Soc. 56: 86, t. 14. (2008). TYPE: Laos, Champasak, Paksong Distr., Bolaven Plateau, Tad
Kamued Waterfall, Picheansoothon & Koontenn 821 (holotype BKF; isotype SING).
4. Monolophus bracteatus (K.Larsen & S.S.Larsen) Veldk. & Mood, comb. nov. -
Caulokaempferia bracteata K.Larsen & S.S.Larsen, Nordic J. Bot. 22; 41 1 , t. 2. (2003). TYPE:
Thailand, Chatuchak Market in Bangkok, originally fromNong Khai, Larsen 4733 7 (holotype
AAU; isotype BKF).
5. Monolophus chayanianus (Tiyaw.) Veldk. & Mood, comb. nov. - Caulokaempferia
chayaniana Tiyaw., Telopea 12: 480, t. 1, 2 (2010). TYPE: Thailand, Mae Hong Son, Pai Distr.,
Doi Jik Jong, Picheansoothon 1017 (holotype BKF; isotypes BK, SING).
6. Monolophus coenobialis Hance, J. Bot. 8: 75 (1870). - Kaempferia coenobialis (Hance)
C.H.Wright, J. Linn. Soc., Bot. 36: 68 (1903). - Caulokaempferia coenobialis (Hance)
K.Larsen, Bot. Tidsskr. 60: 177 (1964). TYPE: China, North River, Fi-loi-tsz Monastery,
Sampson 11369 (holotype K),
Roscoea flava Merr., Lingnan Sci. J. 13: 21 (1934). - Type: China, Kwangtung [Guangdong],
Tsungfa-Lungmoon Distr., Sam Kok Shan, CCC 20595 (W.T. Tsang) (holotype SYS; isotypes
E, possibly in A).
7. Monolophus jirawongsei (Picheans. & Mokkamul) Veldk. & Mood, comb. nov. -
Caulokaempferia jirawongsei Picheans. & Mokkamul, Folia Malaysiana 5: 75, t. 9-15 (2004).
TYPE: Thailand, Nong Khai, Phu Wua, Picheansoothon & Mokkamul 759 (holotype BKF;
isotypes BK, I<:EP, SING).
8. Monolophus khaomaenensis (Picheans. & Mokkamul) Veldk. & Mood, comb. nov. -
Caulokaempferia khaomaenensis Picheans. & Mokkamul, Folia Malaysiana 5: 6, 1 2, 3B, 4- 7.
(2004). TYPE: Thailand, Nakhon Si Thanmiarat, summit of Kliao Maen, Mokkamul 130703-01
(holotype BKF; isotypes BK, KEP, SING).
9. Monolophus kuapii (K.Larsen) Veldk. & Mood, comb. nov. - Caulokaempferia kuapii
K.Larsen, Bot. Tidsskr. 60: 175, t. 5 (1964). TYPE: Thailand, Chantaburi, Mt. Khao Kuap, Put
3016 (holotype C).
1 0. Monolophus larsenii (Suksathan & Triboun) Veldk. & Mood, comb. nov. - Caulokaempferia
larsenii Suksathan & Triboun, Edinburgh J. Bot. 60: 513, t. 1. (2004). TYPE Thailand, Chiang
Mai, Doi Phe Pan Nam, : Suksathan, Triboun & Wongnak 3429 (holotype QBG; isotypes AAU,
BK, BKF, E, K).
11. Monolophus limianus (Mokkamul & Picheans.) Veldk. & Mood, comb. nov. -
Caulokaempferia limiana Mokkamul & Picheans., Folia Malaysiana 5: 188, t. 1,2, 3, 5, 8
(2004). TYPE: Thailand, Phitsanulok ,Chat Trakan Waterfalls National Park, Picheansoonthon
& Mokkamul 742 (holotype BKF; isotypes BK, KEP, SfNG).
12. Monolophus linearis (Wall.) Wall., Numer. List 6592 (1832). - Kaempferia linearis Wall,
in Roxb., FI. Ind. 1: 20 (1820). - Costus linearis (Wall.) Spreng., Syst. Veg. 1: 13 (1824,
Nomenclatural changes in Zingiberaceae
227
“1825”). — Caulokaempferia linearis (Wall.) K.Larsen, Bot. Tidsskr. 60: 170 (1964). TYPE:
India, Silhet, Gomez (“lEG.”) in Wallich 6592 (holotype K; isotypes C, CAL, E, L, P, IDC
microfiche 7394).
13. Monolopitus pedemontanus (Triboun & K.Larsen) Veldk. & Mood, comb. nov. -
Caulokaempferia pedemontana Triboun & K.Larsen in K.Larsen et al., Nordic J. Bot. 23:
403, t. 3 (2005). TYPE: Thailand, Nakon Nayok, Nang Rong Tails, Larsen, Larsen, Tange &
Niyomdham 43774 (holotype AAU; isotype BK).
14. Monolophus petelotii (K.Larsen) Veldk. & Mood, comb. nov. - Caulokaempferia petelotii
K.Larsen, Bot. Tidsskr. 80: 176, t. 6. (1964). TYPE: Vietnam, Chapa, Petelot s.n. Jul 1924
(holotype P).
15. Monolophus phulangkaensis (Picheans.) Veldk. & Mood, comb. nov. - Caulokaempferia
phulangkaensis Picheans., Taiwania 53: 249, t. 3, 4, 7 (2008). TYPE: Thailand, Loei, Phu
Luang, Kok Nok-kraba, Picheansoothon 729 (holotype BKF; isotypes BK, SING).
16. Monolophus phuluangensis (Picheans. & Mokkamul) Veldk. & Mood, comb. nov. -
Caulokaempferia phuluangensis Picheans. & Mokkamul, Folia Malaysiana 5: 70, t. 1, 2, 4-8
(2004). TYPE: Thailand, Loei, Phu Luang, Kok Nok-kraba, Picheansoonthon & Mokkamul
739 (holotype BKF; isotypes BK, KEP, SING).
17. Monolophus phutokensis (Picheans.) Veldk. & Mood, comb. nov. - Caulokaempferia
phutokensis Picheans, in Picheans. & Koonterm, Taiwania 53: 253, t. 5-7. (2008). TYPE:
Thailand, Nong Khai, Amphoe Si Wilai, Phu Tok Noi, Picheansoonthon 732 (holotype BKF;
isotypes BK, SING).
18. Monolophus phuwoaensis (Picheans. & Koonterm) Veldk. & Mood, comb. nov. -
Caulokaempferia phuwoaensis Picheans. & Koontenn, Taiwania 53: 248, t. 1, 2, 7 (2008).
TYPE: Thailand, Nong, Khai, Amphoe Bung Klila, Pu Woa Wildlife Sanctuary, Picheansoothon
723 (holotype BKF; isotypes BK, SING).
19. Monolophus saksuwaniae (K.Larsen) Veldk. & Mood, comb. nov. - Caulokaempferia
saksuwaniae K.Larsen, Bot. Tidsslcr. 68: 157, 1. 1 . (1973). TYPE: Thailand, Phangnga, Takuapa,
Mt. Khao Pra Mi, Larsen, Larsen, Nielsen & Santisuk 30832 (holotype AAU).
20. Monolophus satunensis (Picheans.) Veldk. & Mood, comb. nov. - Caulokaempferia
satunensis Picheans., Folia Malaysiana 8: 56, t. 15 (2007). TYPE: Thailand, Satun, Thung Wa,
Tan Pliu Waterfall, Picheansoonthon 531 (holotype BKF; isotypes KEP, SING).
21 . Monolophus saxicola (K.Larsen) Veldk. & Mood, comb. nov. - Caulokaempferia saxicola
K.Larsen, Bot. Tidsslcr. 60: 171, t. 3, 4 (1964). TYPE: Thailand, Prachinburi, Mt. Khao Khieo
Hills, Larsen s.n. (holotype C).
22. Monolophus secundus (Wall.) Wall., Numer. List 6591 (1 832) {“secimda”). - Kaempferia
secunda Wall, in Roxb., FI. Ind. 1:19 (1820). - Costus secundus (Wall.) Spreng., Syst. Veg. 1:
13 (1824, “1825”). - Caulokaempferia secunda (Wall.) K.Larsen, Bot. Tidsskr. 60: 1 70 (1964).
228
Card. Bull. Singapore 66(2) 2014
TYPE: India, Silhet, F.D. & W.G. (Da Silva and Gomez) in Wallich 6591 (holotype K; isotypes
C, CAL, P, IDC microfiche 7394).
23. Monolophus sikkimensis (King ex Baker) Veldk. & Mood, comb. nov. - Kaempferia
sikkimensis King ex Baker in Hook.f., FI. Brit. India 6: 223 (1890). - Caulokaempferia
sikkimensis (King ex Baker) K.Larsen, Bot. Tidsskr. 60: 169 (1964). TYPE: India, Sikkim,
Elwes s.n., Aug 1886 (lectotype K, designated here).
24. Monolophus sirirugsae (Ngamr.) Veldk. & Mood, comb. nov. - Caulokaempferia
sirirugsae Ngamr., Nordic J. Bot. 26: 325, t. 1, 2 (2009). TYPE: Thailand, Phangnga, Khao
Lumpee Waterfall, Ngamriahsakul 51-01 (holotype BKF; isotypes PSU, Wailak Univ. Herb.).
25. Monolophus tamdaoensis (Picheans. & Inthar.) Veldk. & Mood, comb. nov. -
Caulokaempferia tamdaoensis Picheans. & Inthar., J. Jap. Bot. 89: 134 (2014). TYPE:
Picheansoonthon & Phokham 180812-1 (holotype BK).
Appendix 2. Taxa excluded from Monolophus (name in bold is currently accepted name).
1. Gastrochilus ochroleucus Ridl., J. Straits Branch Roy. Asiat. Soc. 32: 109, 110. (1899)
{"^ochroleucd’"). - Kaempferia ochroleuca (Ridl.) K.Schum. in Engler, Pflanzenr. IV, 46, Heft
20: 75. (1904). - Boesenbergia ochroleuca (Ridl.) Schltr. in Fedde, Repert. Spec. Nov. Regni
Veg. 12: 316. (1913). TYPE: Thailand, between Kasum and Pungah, Ridley Illustration 00754
(lectotype SING, designated by Turner 2000: 25-26, t. 2; isotype K?).
Note - Included in Gastrochilus Wall, non D. Don unranked § Mesanthi by Ridley (1899: 109)
and in Kaempferia subg. Monolophus by Schumann (1904).
2. Kaempferia L. subg. Pyrgophyllum Gagnep., Bull. Soc. Bot. France 48: Ixxviii (1902,
“1901”). — Camptandra Ridl. sect. Pyrgophyllum (Gagnep.) Gagnep. ex K.Schum. in Engler,
Pflanzenr. IV, 46, Heft 20: 63 (1904). - Caulokaempferia K.Larsen sect. Pyrgophyllum
(Gagnep.) R.M.Sm., Notes Roy. Bot. Gard. Edinburgh 31: 291 (1972). - Pyrgophyllum
(Gagnep.) T.L.Wu & Z.Y.Chen, Acta Phytotax. Sin. 27: 126 (1989). TYPE: Kaempferia
yunnanensis Gagnep.
Note - Pyigophyllum in our analysis (Fig. 1) is distinct from Monolophus (see below #10).
3. Kaempferia decus-sylvae Hallier f., Ann. Jard. Bot. Buitenzorg 13: 321, t. 27, f 4. (1896).
- Haplochorema decus-sylvae (Hallier f.) Valeton, Bull. Jard. Bot. Buitenzorg ser. II, 27: 116.
(1918). TYPE: Indonesia, Kalunantan, Liang Gagang Mts, Hallier 2636 (lectotype L [122322
spirit], designated here; isotype BO).
Note - Attributed to Kaempferia sect. Monolophus by Hallier.
4. Kaempferia elegans Wall., PI. Asiat. Rar. 24, t. 27 (1829). - Monolophus elegans (Wall.)
Wall., Numer. List 6593 (1832); Horan., Prodr. Monogr. Scitam.: 22 (1862). TYPE: Wallich
(1829) cited “Pegu, Martaban, Rangoon, Amherst, Moalmyne”. Only the Rangoon locality is
mentioned in the Wallich Herbarium, K-W. We here select 6593-A, bottom specimen-{\Qcto\ypQ
K-W [IDC microfiche 7394], first step designation by Picheansoonthon & Koonterm (2008b:
77, “Tenasserim”), second step designated here: Bimia, Rangoon.
Note - Included in Kaempferia subg. Monolophus by Baker (1892).
Nomenclatural changes in Zingiberaceae
229
5. Kaempferia gracillima K.Schuin. in Engler, Pflanzenr. IV, 46, Heft 20: 74 (1904). -
Camptandra gracillima (K.Schum.) Valeton, Bull. Jard. Bot. Buitenzorg II, 27: 115. (1918).
TYPE: Sarawak, Selebut, Haviland 448 (holotype B, lost; isotype K). = Camptandra parvula
(King ex Baker) Ridl. van angustifoUa Ridl,, fide Newman et al. (2004: 61).
Note — Included in Kaempferia subg. Monolophus by Schumann (1904: 74).
6. Kaempferia macrochlamys Baker in Hook.f., FI. Brit. India 6: 223. 1892. - Stahlianthus
macrochlamys (Baker) Craib, Bull. Misc. Inform. Kew. 1912: 401 (1912). TYPE: Burma,
Tenasserim, Parish s.n. (holotype K).
Note -Included in Kaempferia subg. Monolophus by Baker (1892).
7. Kaempferia pai^ula King ex Baker in Hook.f., FI. Brit. India 6: 223 (1892). - Camptandra
parvula (King ex Baker) Ridl., J. Straits Branch Roy. Asiat. Soc. 32: 104. (1 899). TYPE: Malay
Peninsula, Goping, King’s collector (holotype K [000255350]; isotype CAL, K).
Note - Included in Kaempferia subg. Monolophus by Baker.
8. Kaempferia philippinensis Merr., Philipp. J. Sci., sect. C, Bot. 10: 296 (1915). TYPE:
Philippines, Luzon, Laguna, San Antonio, Dahican River, Ramos BS 14952 (holotype PNH,
lost).
Note - Included m Kaempferia subg. Monolophus by Loesener (1 930: 566) and Merrill (1934).
9. Kaempferia siphonantha King ex Baker in Hook.f, FI. Brit. India 6: 222 (1892). -
Boesenbergia siphonantha (King ex Baker) M.Sabu et al., Rheedea 14: 55, t.l, 2. 2004.
TYPE: Andaman Isl, King’s collector 372 (probably Kunstler) (holotype K [000640517];
isotype CAL).
Note - Included in Kaempferia subg. Monolophus by Baker (1892).
10. Kaempferia yunnanens is Gagnep., Bull. Soc. Bot. France 48: Ixxvii (1901). - Camptandra
yunnanensis (Gagnep.) K.Schum. in Engler, Pflanzenr. IV, 46, Heft 20: 64. (1904). —
Caulokaempferia yunnanensis (Gagnep.) R.M.Sm., Notes Bot. Gard. Edinburgh 31: 292,
t. 2. (1972). — Monolophus yunnanensis (Gagnep.) T.L.Wu & S.J.Chen in Anon., Iconogr.
Connophyt. Sin. 5: 586, t. 8001 (1976), nom. inval,, sine basion.; T.L.Wu & S.J.Chen
(“Senjen”) Acta Phytotax. Sin. 16(3): 29 (1978). - Pyrgophyllum yunnanensis (Gagnep.)
T.L.Wu & Z.Y.Chen, Acta Phytotax. Sin. 27: 127 (1989). TYPE: China, Yunnan, N of Tali,
Tsong-so gorges (Ten tchouan), Delavay 2721 (holotype P; isotype E).
Kaempferia fongyuensis Gagnep., Bull. Soc. Bot. France 48: Ixxviii (1901). - Camptandra
fongyuensis (Gagnep.) K.Schum. in Engler, Pflanzenr. IV, 46, Heft 20: 64. (1904). TYPE:
China, Yunnan, Fung- Yu, near Er’Hai lake, d ’Orleans s.n., 19 Jun (holotype P) (synonymy
fide Wu & Chen, 1978: 29).
1 1 . Monolophus philippianus A.Dietr. , Allg. Gartenz. 1 7 : 266 ( 1 849). - Kaempferia philippiana
(A. Dietr.) K.Schum. in Engler, Pflanzenr. lV,46,Heft20: 85 {\90A).-Stahlianthusphilippianus
(A.Dietr.) Loes. in Engler, Nat. Pflanzenfam., ed. 2, 15a: 564 (1930). TYPE: From India, sent to
Remecke of the Decker Garten, Berlin, flowered in 1849, Philippi A° 1 846 (holotype B, lost).
12. Monolophus scaposus (Nimmo) Dalzell in Hook., J. Bot. Kew Gard. Misc. 2: 143 (1850). -
Hedychium scaposum Nimmo in Graham, Cat. PI. Bombay: 205 (1839). —Kaempferia scaposa
(Nimmo) Benth. in Benth. & Hook.f, Gen. PI. 3: 642 (1883). - Curcuma scaposa (Nimmo)
230
Card. Bull. Singapore 66(2) 2014
Skomick. & M.Sabu, Ann. Bot. (Oxford) 100: 524 (2007). TYPE: India, southern Concan,
v
Kings.}!, (neotype BM, designated by Leong-Skomickova et al. (2007); isoneotype K).
Note - Included in Kaempferia L. sect. Stachyanthesis Benth. (1883). According to Leong-
V
Skomickova et al. (2007) Hedychium scaposmn Nimmo is the basionyin of Monolophus
scaposus even though the name was not mentioned by Dalzell (1850). Later, Dalzell & Gibson
(1861) described Hedychium scaposum Nimmo with M. scaposus Dalz. in synonymy. Under
Art 41.4 of the ICN, even though there is no reference to a basionyin, DalzelTs intention is
obvious and Nimmo is to be accepted as the author of the basionyin. As no Graham nor Nimmo
material apparently exists Leong-Skomickova et al. (2007) designated a neotype.
Appendix 3. New Boesenbergia combinations.
Boesenbergia Kuntze, Revis. Gen. PI. 2 (1891) 685. - Gastrochilus Wall., PI. Asiat. Rar. 1
(1829) 22. TYPE: Boesenbeigia pulcheirima (Wall.) Kuntze, lectotype designated by Holttum
(1950: 107).
Jirawongsea Picheans., Folia Malaysiana 9:2 (2008). TYPE: Jirawongsea laotica (Picheans.
& Mokkamul) Picheans.
1. Boesenbergia alba (K.Larsen & R.M.Sm.) Mood & L.M.Prince, comb. nov. -
Caidokaempferia alba K.Larsen & R.M.Sm., Notes Roy. Bot. Gard. Edinburgh 31: 288, t. 1
(1972). - Jirawongsea alba (K.Larsen & R.M.Sm.) Picheans., Folia Malaysiana 9: 3, t. 1^.
(2008). TYPE: Thailand, Phitsanulok, summit Phu Mieng, Smitinand & Wd/-ncke 949 (holotype
AAU; isotype BKF).
2. Boesenbergia burttii (K.Larsen & Jenjitt.) Mood & L.M.Prince, comb. nov. - Caidokaempferia
burttii K.Larsen & Jenjitt., Edinburgh J. Bot. 60: 509, t. 1. (2004). - Jirawongsea burttii
(K.Larsen & Jenjitt.) Picheans, Folia Malaysiana 9: 3, t. 5-8 (2008). TYPE: Laos, Champassak,
Paksong (Bolaven Plateau), Jenjittikul 8263 (holotype AAU; isotypes BKF, E, PBM).
3. Boesenbergia laotica (Picheans. & Moklcamul) Mood & L.M.Prince, comb. nov. -
Caulokaempferia laotica Picheans. & Mokkamul, Nat. Hist. Bull. Siam Soc. 54: 75, t. 1-5.
(2006). - Jimwongsea laotica (Picheans. & Mokkamul) Picheans., Folia Malaysiana 9: 4, t.
7, 8 (2008). TYPE: Laos, Champasak, Phu Luang, Bolaven Plateau, Picheansoonthon 581
(holotype BKF; isotype SING),
4. Boesenbergia thailandica (K.Larsen) Mood & L.M.Prince, comb. nov. - Caulokaempferia
thailandica K.Larsen, Bot. Tidsskr. 68: 157, t. 2. (1973). TYPE: Thailand, Loei, Phu Kradung,
Smitinand 336 (holotype BKF).
5. Boesenbergia violacea (K.Larsen & Triboun) Mood & L.M.Prince, comb. nov. -
Caulokaempferia violacea K.Larsen & Triboun in K.Larsen, Nordic J. Bot. 22: 414, t. 3, 4.
(2003). TYPE: Thailand, Loei, Phu Rhua, Triboun 399 (holotype BK; isotypes AAU, BKF).
Appendix 4: GenBank accession numbers for Zingiberaceae used in this study. Format:
identification, plant sample number [for newly sequenced samples] (collector and voucher
number, herbarium), ITS GenBank number/rmif GenBank number.
Nomenclatural changes in Zingiberaceae
231
Boesenbergia alba (K.Larsen & R.M.Sm.) Mood & L.M.Prince: M3253 {Mood 3253, BISH)
KF982792/KF982801.
Boesenbergia alba (K.Larsen & R.M.Sm.) Mood & L.M.Prince: M12C30 {Funakoshi s.n.,
MBK) KF982793/KF982802.
Boesenbergia bnrttii (K.Larsen & Jenjitt.) Mood & L.M.Prince: M12C32 (Funakoshi s.n.,
MBK) KF982794/KF982803.
Boesenbergia curtisii (Baker) Schltn: Ml 739 {Mood 1739, BISH) KF982795/KF982804
Boesenbergia hamiltonii Mood, S. Dey & L.M.Prince: M301 7 {Mood 301 7, BISH) JX992754/
JX992815.
Boesenbergia plicata van lurida Holttum: M3 120 {Mood & Vatcharakorn 3120, BK)
JX992808/JX992839.
Boesenbergia pulcherrima (Wall.) Kuntze: M08P276 {Mood 08P276, BISH) X992809/
JX992748.
Boesenbergia rotunda (L.) Mansf.: {Kress 00-6737, US) AF478726/AF478726.
Boesenbergia thailandica (K.Larsen) Mood & L.M.Prince: {Ngamriabsakul 61, BKF)
AY424748/-.
Boesenbergia violacea (K.Larsen & Triboun) Mood & L.M.Prince: M12C31 {Funakoshi s.n.,
MBK) KF982796/KF982805.
Camptandra paimda (King ex Baker) Ridley: {Kress 99-6328, US) AF478730/AF478830.
Cornukaempferia aurantiflora Mood & K.Larsen: {Kress 01-6983, US) AF478736/AF478835.
Distich ochlamys rubrostriata W. J.Kress & Rhese: {Kress 01-6848, US) AF478745/AF478844.
Globba curtisii Holttum: {Kress 99-6347, US) AF478754/AF478853.
Haniffia albiflora K.Larsen & Mood: (Kress 99-6370, US) AF478756/AF478855.
Hedychiunt bordelonianum W.J.Kress & K.J. Williams: {Kress 99-6462, US) AF478757/
AF478856.
Herniorchis rhodorrhachis K.Schum.: {Newman 861, E) AF478763/ AF478863.
Hitchenia glauca Wall.: {Kress 00-6743 US) AF478765/AF478864.
Hornstedtia hainanensis T.L.Wu & S.J.Chen: {Kress 97-5769, US) AF478766/AF478865.
Kaempferia rotunda L.: {Kress 99-6304, US) AF478767/AF478868.
Monolopltus coenobialis Hance: (unknown) HM236121/".
Monoloplius limianus (Mokkamul & Picheans.) Veldk. & Mood.: M3246 {Mood 3246, BISH)
KF982797/KF982806.
Monolopltus linearis (Wall.) Wall.: M12C193 (Dey NU198, CAL) KF982798/KF982807.
Monoloplius saxicola (K.Larsen) Veldk. & Mood: M12C29 {Funakoshi s.n., MBK) KF982799/
KF982808.
Monoloplius sikkimensis (King ex Baker) Veldk. & Mood: New2458 {Newman 2458, spirit,
E) KF982800/KF982809.
Pommereschea lackneri Witt.: {Kress 00-6739, US) AF478776/AF478877.
Pyrgophyllum yunnanense (Gagnep.) T.L.Wu & Z.Y.Chen: {Kress 00-6596, US) AF478777/
AF478878.
Rhyncliantlius beesianus W.WSm.: {Kress 97-5827, US) AF478784/AF478885.
Roscoea purpurea Sm.: {Kress 01-6953, US) AF478787/AF478888.
Siliquamomum tonkinense Baill.: {Kress 00-6802, US) AF478791/AF478892.
Siphonochilus kirkii (Hook.f.) B.L.Burtt: {Kress 94-3692, US) AF478794/AF478895.
Tamijia flagellaris S.Sakai & Nagam.: {Kazuyuki S55, KYOTO) AF478797/AF478898.
Zingiber gramineum Noronha ex Blume: {Kress 96-5739, US) AF478800/AF478902.
Gardens’ Bulletin Singapore 66(2): 233-239. 2014
233
Conservation status and lectotypfication of Alangium
ridleyi (Cornaceae) in Singapore
L. Wijedasa^’^’^ Z.Q. Shee^ & E. Chia^
’Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569
^Applied Entomology Lab, Department of Biological Sciences,
National University of Singapore, 14 Science Drive 4, Singapore 117543
^Rimba, 18E Kampung Basung, Kuala Berang 21700,
Terengganu, Malaysia
lahirux@gmail .com
ABSTRACT. Alangium ridleyi King is lectotypified and the conservation status updated from
Nationally Extinct to Endangered in Singapore and Endangered in Penmsula Malaysia.
Keywords. Alangium ridleyi King, Endangered, lectotypification. Peninsula Malaysia,
Singapore
Introduction
The genus Alangium Lam. in the family Cornaceae comprises 24 species of woody
trees and climbers distributed in the tropical and subtropical regions of the Old World
(Berhaman, 1995; Feng et ah, 2009). Five species oi Alangium have been recorded in
Singapore, of which one, Alangium ndleyi King, has been presumed to be nationally
extinct in Singapore (Tan et ah, 2008; Chong et ah, 2009).
Based on observations of the population of this species in the Singapore Botanic
Gardens, H.N. Ridley, Director of Singapore Botanic Gardens (SBG) from 1888 to
1912, believed this tree could be a new species. When the trees eventually did flower
in 1 892, Ridley collected specimens and sent them to his colleague, Sir George King,
the superintendent at the Royal Botanic Garden, Calcutta, who subsequently described
the species as Alangium ridleyi in 1902 in honour of Ridley. The flowering in 1892
was documented by Ridley and the Gardens’ botanical artists of the time, J.B. de Alwis
and C.G. de Alwis. The painting is deposited in the SING library (Fig. 1).
There is some confusion on the exact type locality of this tree in the Gardens. In
the Flora of the Malay Peninsula, Ridley ( 1 922) stated the following: ‘I have only seen
one tree. Singapore, rockery in Gardens.’ The rockery, during Ridley’s time, was the
location of the current Fernery, which is adjacent to the Gardens Jungle and consisted
of trees which originally belonged to the Gardens Jungle. While King’s description
includes specimens from Singapore and the Malaya Peninsula, he lists the distribution
as ‘SINGAPORE, m the Botanic Garden Jungle, Ridley 4941’ with no mention on
the number of sheets nor the herbaria in which the specimens were deposited, as such
all material are syntype material. We lectotypify the species based on a study of the
234
Card. Bull Singapore 66(2) 2014
Fig. 1. The 1892 painting of Alangium ridleyi King by J. De Alwis, H.N. Ridley and C. De
Alwis. Reproduced with permission from the Singapore Botanic Gardens.
Alangium ridleyi in Singapore
235
available material at SING, CAL and K. We also assess the conservation status of
Alangium ridleyi in Singapore and Peninsular Malaysia.
A recent prolonged dry spell in Singapore, which lasted for a period of nine
weeks from mid-January 2014 through to mid March 2014 saw new records for the
driest month since records began in 1 869 as well as the windiest month in the last 30
years (National Environment Agency, 2014). Only 0.2 mm of rain were recorded in
February 2014, compared to average monthly rainfalls of 161 mm since 1869. Since
the dry spell, many species of plants in various habitats throughout Singapore have
been observed to flower, including the Alangium ridleyi mentioned here (Fig. 2). This
has stimulated new interest in this species.
Taxonomic notes
Alangium ridleyi King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 71(1): 78. (1902);
Ridley, FI. Malay Penin. 1: 893 (1922). TYPE: Singapore Botanic Gardens, 1892,
H.N. Ridley 4941 (lectotype SING! [SING0059170], designated here; isolectotypes
CAL!, K! [K00007704].
Conservation Status
To assess the conservation status of Alangium ridleyi we used herbarium specimen
distribution data fi'om SING and KEP (Fig. 3) for Peninsular Malaysia and Singapore
to calculate the extent of occurrence (EOO) and area of occupancy (AGO) using
GeoCAT (Baclunan et al., 2011) - a software used to generate lUCN Red List
assessments (lUCN, 2001) using distribution area alone. While a Global Biodiversity
Inventory Facility (2014) search revealed the potential presence of this species in
Indonesia, Thailand and Vietnam, this could not be verified. As such the conservation
assessments are national ones for Singapore and Malaysia where identity of the
specimens is confirmed.
Our assessment of the lUCN conservation status of Alangium ridleyi in
Peninsular Malaysia assessed by GeoCAT was Endangered (EN B2ab(iii)) based on an
estimated area of occupancy (AOO) of less than 100 km^ (using the lUCN default cell
width of 2 km). Of the 32 documented specimen localities, eight (i.e. 25%) are within
protected areas while a further fifteen (46%) appear to be in forested areas outside of
protected areas and subject to disturbance and decline.
In Singapore, the latest version of the Singapore Red Data Book (Davison et
al., 2008) and the checklist of vascular flora for Singapore (Chong et al, 2009) list the
conservation status of this species as Nationally Extinct. However, remnants of the
original population of trees in the Botanic Gardens stil 1 exists. In addition two additional
trees, one in MacRitchie reservoir and another in Mandai forest, have recently been
observed although they lack specimens for verification. This is a reduction from the
236
Card. Bull Singapore 66(2) 2014
Fig. 2. Alangium ridleyi King. A. Flowering branch. B. Close up of flowers. Photographed at
Singapore Botanic Gardens, Lawn H, Ace. No. 00/705 1*A. (Photo: Edmund Chia)
Alangium ridleyi in Singapore
237
Pulau Pinanbpii^ PjAa
Kelantan
SOUTH
CHINA
SEA
PENINSULAR
MALAYSIA
Pahang
Selaaafir
Negeri Sembllan
Melaka
SUMATRA
200
Kilometers
Fig. 3. Distribution of Alangium ridleyi King in Peninsular Malaysia and Singapore.
(Distribution data are from SfNG and KEP).
former distribution based on specimen data (Table 1). Our conservation assessment
for Singapore is Critically Endangered (CR D) based on the very small number of
remaining individuals in Singapore.
238
Card. Bull. Singapore 66(2) 2014
Table 1. Previous collections of Alangium ridleyi in Singapore in herbaria.
Collector
Herbarium
Collector
Number
Date
Collected
Locality
Samsuri, A.
SING
158
29 July 2003
Nee Soon Swamp Eorest
Tang, E.
SING
393
27 March 1995
SBG
Ngadiman, I.
SING, KEP
36136
23 April 1940
Bkt. Timah
Kiah, S.
SING
s.n.
20 August 1940
Bkt. Mandai
Ridley, H.N.
SING,K
4941
1893
SBG, Rain Forest
Cantley, N.
SING
s.n.
1880
ACKNOWLEDGEMENTS. We thank Dr David Middleton and two anonymous reviewers for
conuTients on the manuscript. Dr Kaliyamurthy Karthigeyan for assistance in viewing the type
specimens at CAL. Dr Richard Chung for sharing the distribution data of specimens in KEP.
Sunia Teo, Craig Williams, Ali Ibrahim and Lua Hock Keong for information on potential
populations of Alangium ridleyi in MacRitchie and Mandai.
References
Bachman, S., Moat, J., Hill, A.W., de la Torre, J. & Scott, B. (2011). Supporting Red List threat
assessments with GeoCAT: Geospatial Conservation Assessment Tool. In: Smith, V. &
Penev, L. (eds) e-Infrastructures for data publishing in biodiversity science. ZooKeys
150:117-126.
Berhaman, A. (1995). Alangiaceae. In: Soepadmo, E. & Wong, K.M, (eds) Tree Flora of Sabah
and Sarawak 1 : 5-1 7. Kuala Lumpur: Ampang Press Sdn. Bhd.
Chong, K. Y., Tan, H.T. W. & Corlett, R.T. (2009). A Checklist of the Total Vascular Plant
Flora of Singapore: Native, Naturalised and Cultivated Species. Raffles Museum of
Biodiversity Research, National University of Singapore, Singapore. 273 pp. http://
nnbr.nus.edu.sg/raffIes_museum_pub/flora_of_singapore_tc.pdf
(accessed on 14 Apr. 2014).
Davison, G.W.H., Ng, P.K.L. & Ho, H.C. (eds) The Singapore Red Data Book: Threatened
plants and animals of Singapore, 2'“^ ed. Pp. 285. Singapore: The Nature Society
(Singapore).
Feng, C.M., Manchester, S.R., & Xiang, Q.Y.J. (2009). Phytogeny and biogeography of
Alangiaceae (Comales) inferred from DNA sequences, morphology, and fossils. Molec.
Phylogenet. EvoL 51(2): 201-214.
Global Biodiversity Inventory Facility (2014). http://data.gbif org/ (accessed on 9 Oct. 2014).
lUCN (2001). lUCN Red List Categories and Criteria: Version 3.1. 2nd ed. Switzerland, Gland
and UK, Cambridge: lUCN.
National Enviromiient Agency (2014). Dry Spell Advisory (4 Mar). Joint Advisory between
NEA and PUB. http://app2.nea.gov.sg/corporate-flinctions/newsroom/advisories/
year/20 1 4/month/3/category/weather-climate/dry-spell-advisory-(4-mar) (accessed
onl2 Aug. 2014).
Alangium ridleyi in Singapore
239
Ridley, H.N. (1922). The Flora of the Malay Peninsula 1: 893. London: L. Reeve & C.o Ltd.
Tan, H.T.W., Tan, K.-X., Ali bin Ibrahim, Chew, RT., Chua, K.S., Duistermaat, H., Ganesan,
S.K., Goh, M.W.K., Gwee, A.T., Kiew, R., Lee, S.M.L., Leong, R, Lim, J., Lok, A.F.S.L.,
Loo, A.H.B., Lum, S.K.Y., Morgany, T., SaiRiddin bin Suran, Sim, S., Haji Samsuri bin
Haji Ahmad, Wee, Y.C., Yap, K.F., Yeo, C.K. & Yong, J.W.H. (2008). Checklists of
threatened species — Seed plants. In: Davison, G.W.H., Ng, R.K.L. & Ho, H.C. (eds) The
Singapore Red Data Book: threatened plants & animals of Singapore, Edition. Rp.
213-244. Singapore: The Nature Society (Singapore).
Turner, I.M. (1997). A catalogue of the vascular plants of Malaya. Gard. Bull. Singapore
47(1): 1-757.
Gardens’ Bulletin Singapore 66(2): 241-242. 2014
241
BOOK REVIEW: Flora of Peninsular Malaysia. Series II: Seed Plants, Volume 4.
(Malayan Forest Records No. 49) R. Kiew, R.C.K. Chung, L.G. Saw & E. Soepadmo
(eds). 2013.
Kepong: Forest Research Institute Malaysia. 25.7 cm x 18 cm, hard cover. 405 pp. ISBN 978-
967-0622-08-8. Price RM 100 / US$ 75.
The Flora of Peninsular Malaysia project is
progressing remarkably well, with the fourth volume
of the seed plants series released late last year. This
volume includes eight taxonomic revisions, namely
Actinidiaceae, Cabombaceae, Crypteroniaceae,
Goodeniaceae, Meliaceae, Memecylaceae,
Opiliaceae and Pandaceae, altogether 166 taxa in 29
genera. As anticipated, this volume is well structured
and produced.
The volume begins with a short chapter on
conservation by L.S.L. Chua that includes two tables
summarising (i) the number of native taxa for each
Red List category assessed using the lUCN Red List
Categories and Criteria (Version 3.1), and (ii) a list of
threatened (i.e.. Critically Endangered, Endangered
and Vulnerable), Data Deficient and Rare taxa, recorded in the volume. The taxonomic
accounts of the eight families are then presented in alphabetical order. The format of
each revision is quite standard, as in any long-running Flora series, following styles
adopted for the first volume. There are taxonomic keys provided to distinguish genera,
as well as species, for all the families revised, and these keys are easy-to-use.
The bulk of this volume is the account of the Meliaceae by D.J. Mabberley and
C.M. Pannell, with a contribution by J.M. Edmonds for Toona. The account has a total
of 15 genera and 106 species of which 51 species are in the genus Aglaia. This genus
was revised by Caroline M. Pannell who has spent many years working diligently
on this very diverse genus and who has already contributed taxonomic accounts for
the Tree Flora of Malaya (1989), Flora Malesiana (1995), and the Tree Flora of
Sabah and Sarawak (2007). Also included in the Meliaceae account is a list of useful
spot characters to identify Meliaceae genera in Peninsular Malaysia. This list is not
only useful for botanists, but also foresters and naturalists. A new term ‘berryoid’ is
introduced to define the fruit type of some Meliaceous taxa, such as Aglaia, Lansium,
Pseudoclausena, Reinwardtiodendron and Walsura, that cannot be considered either a
berry or a capsule based on the current classification of fruit types.
The second largest family in the volume is the Memecylaceae which includes
the two gQnQra Lijndenia (1 species) mdMemecylon (32 species). The well-researched
account by M. Hughes contains some very informative figures to guide users through the
taxonomy of Peninsular Malaysian Memecylon based on the morphological characters
of the anther, if fertile materials are available, or based on leaf size, shape and venation
prominence, when only non-fertile materials are available. However, I noticed that in
242
Card. Bull. Singapore 66(2) 2014
Figure 3 (page 278), an illustration of “M megacarpunf' is included amongst a selection
of Peninsular Malaysian Memecylon species with distinct marginal veins. Perhaps it
would have been better to illustrate “M. pseudomegacarpum’' instead ofM megacarpum
which is endemic to Borneo. Likewise, in die “Foreword” and “Acknowledgements”
sections, the family is erroneously refen*ed to as “Memecylonaceae”.
In contrast to these two large-sized families, the rest are fairly small and
straightforward, namely Actinidiaceae (2 genera, 11 species), Cabombaceae (1 genus,
2 species), Crypteroniaceae (2 genera, 3 species), Goodeniaceae (1 genus, 1 species),
Opiliaceae (4 genera, 4 species) and Pandaceae (2 genera, 4 species).
This volume is well illustrated with 67 line-drawings representing slightly fewer
than half of the taxa enumerated in the volume. On the other hand, distribution maps
are provided for almost all of the species treated in the volume. At the end, just before
the index, there are 30 pages of colour plates for some of the taxa. With the publication
of this volume, 46 seed plants families (see inner back cover of the book) have now
been revised for the Flora of Peninsular Malaysia and we eagerly look forward to the
next one.
Y.W. Low
Singapore Botanic Gardens
Gardens’ Bulletin Singapore 66(2): 243-244. 2014
243
BOOK REVIEW: A Guide to the Native Palms of Singapore, H.B. Loo, Ang
Wee Foong, William J. Baker & Hugh T.W Tan. 2014.
Singapore: Science Centre. 15 cm x 10.4 cm, card cover. 176 pp. ISBN 978-981-07-8878-0.
Price SGD 7.90.
What a fine little book! Diminutive in size (fits easily
into the pocket of a bush jacket, although that really
is not required in much of Singapore!), concise in
its writing, and well-conceived in its coverage, this
Guide must do more than any other in the same time
spent perusing it.
Singapore is an island nation and it can be
quite astounding how rich its natural vegetation
is, despite the intensive waves of change it has
weathered. For some time now, the most ardent
of Singaporean naturalists have been actively
promoting the value of the native fiora and joining
them today are an entire community of nature lovers
very much into understanding more of what this
inherently rich island has. And now we have our
handy guide to Singapore palms. Palms do not fail
to captivate because of their unique form, yet there is a huge diversity with over 2500
species worldwide.
Palms are generally thought of as large plants; a botanist who collects from a tall
tree palm has few means of obtaining truly representative specimens and in time takes
home nothing short of trophies. The spiny palms, especially the climbing ones called
rotan in Malay (‘rattans’), some of which drape the most majestic of rainforest trees
even, have attracted only the bravest and most stubborn of students. No doubt, the
universal appeal of palms has stemmed from displays in botanical gardens, augmented
by some use in landscaping and street planting. The myriad ways in which palms are
useful to tropical communities are legend. The palms contributing much public appeal
are those that thrive in open, exposed sites and which are therefore more amenable to
cultivation. Thus, building up an interesting diversity of such palms brings with it both
awe and envy because of the increasingly demanding space requirements.
But the other side of the challenge is much less visible or understood. In the
rainforest regions of the world, the understorey palms abound, many of which can
only survive with sufficient shade, humidity and particular soil conditions. And many
must be disappearing with forest conversion and fragmentation. Conserving such
species is always best achieved with their natural habitat, although few public gardens
have ventured into presenting forest understorey palms as a conservation exhibit, or
developing conservation collections to any meaningful extent. There are a number of
smaller pot palms that have found their way from the shade of the rain forest, but the
true diversity of form and elegance in forest palms is a remarkable one.
A Qufd'i ta
244
Gard. Bull. Singapore 66(2) 2014
This Guide is a good and handy primer. The first 39 figures are numbered
for easy reference in the general chapters (Introduction, Economic Importance of
Palms, Palm Characteristics, Why Study Palms in Singapore?) but not the rest of the
illustrations which, placed with their respective taxa, are still easily referred to. Table
5 lists the 54 species of native Singapore palms, with 12 thought to be locally extinct.
The species are introduced via three groupings: the feather (pinnate-leaved) palms, fan
palms, and mostly climbing, scaly-fruited rattan palms, their distinctive fomi and habit
constituting the most logical presentation. The book is the 47th in the Singapore series
sustained so well by the Science Centre Singapore with the support of BP Singapore
Pte Ltd.
K.M. Wong
Singapore Botanic Gardens
The Gardens’ Bulletin
Singapore
VOL. 66 2014
ISSN 0374-7859
VOLUME 66
No. 1
No. 2
pages 1-120 (printed version published 10 July 2014)
pages 121-244 (printed version published 20 November 2014)
THE GARDENS’ BULLETIN, SINGAPORE
The Gardens* Bulletin, Singapore is a peer-reviewed journal publishing original papers and
reviews on a wide range of subjects: plant taxonomy (including revisions), phytogeography,
floristics, morphology, anatomy, as well as horticulture and related fields, such as ecology and
conservation, with emphasis on the plant life of the Southeast Asian-Pacific region.
Dr David J. Middleton Dr Jana Leong-Skomickova
(Editor-in-Chief) (Managing Editor)
S. Lee Y.W. Low Christina Soh
(Graphics Editor) (Copy Editor) (Business Manager)
Editorial Advisory Board
Professor Sir Peter Crane
Yale University
U.S.A.
Dr Rogier PJ. de Kok
Royal Botanic Gardens, Kew
U.K.
Dr W. John Kress
National Museum of Natural History
Smithsonian Institution
U.S.A.
Dr Mark Hughes
Royal Botanic Garden Edinburgh
U.K.
Dr Kiat W. Tan
Gardens By The Bay
Singapore
Dr Nigel P Taylor
Singapore Botanic Gardens
National Parks Board, Singapore
Dr Ian M. Turner
Singapore Botanic Gardens
National Parks Board, Singapore
Dr Jan-Frits Veldkamp
National Biodiversity Center
The Netherlands
Dr Jun Wen
National Museum of Natural History
Smithsonian Institution
U.S.A.
Professor Nianhe Xia
South China Institute of Botany
PR. China
Published twice yearly by the National Parks Board, Singapore, the annual subscription for the journal
is Singapore $100.00 including postage. Overseas subscribers should make payment in the forai of
bank draft or international money order in Singapore cuiTency, payable to National Parks Board. Please
forward payment to “Accounts Receivable Section, National Parks Board, Headquarters,
Singapore Botanic Gardens, 1 Cluny Road, Singapore 259569”.
Copyright ©
National Parks Board
Singapore Botanic Gardens
1 Cluny Road
Singapore 259569
Printed by Oxford Graphic Printers Pte Ltd
CONTENTS OF VOLUME 66 (2014)
VOL. 66 (1), July 2014
D.J. Middleton
Editorial 1
K.H. Lau
The conservation of Peninsular Malaysian Geostachys (Zingiberaceae) 3
J. Leong-Skornickova
Orchidantha lengguanii (Lowiaceae), a new species
from Peninsular Malaysia, and typification of O. maxillarioides 15
J. Leong-Skornickova, H.-J. Tillich & Q.B. Nguyen
Two new species and one new variety of Aspidistra
(Asparagaceae: Nolinoideae) from southern Vietnam 27
J. Leong-Skornickova, H.D. Tran, Q.B. Nguyen & O. Sida
Siliquamomum alcicorne (Zingiberaceae: Alpinioideae),
a new species from central Vietnam 39
Y.W. Low, P.K.F. Leong, S.P. Tee, Rajesh Singh,
M.L.C. Tay & K.M. Wong
Margaritaria (Phyllanthaceae), a new generic record
for the Singapore flora 47
D.J. Middleton
A new combination in Liebigia (Gesneriaceae) 57
T.K.T. Nguyen & N.T. Nguyen
Anew species of Mallotus (Euphorbiaceae) from Vietnam 61
V.D. Nguyen, H.Q. Bui & J. Bogner
The status of Cryptocoryne annamica
(Araceae: Aroideae: Cryptocoryneae) in Vietnam 67
S.M. Scott & D.J. Middleton
A revision of Ornithoboea (Gesneriaceae) 73
VOL. 66 (2), November 2014
Y.F. Deng
Phlogacanthus magnus, a new combination in the Acanthaceae
from Malaysia 121
R. Kiew
Two new species and one new subspecies of Ridleyandra (Gesneriaceae)
from Peninsular Malaysia 125
R. Kiew
Two new white-flowered Codonoboea species (Gesneriaceae)
from Peninsular Malaysia 137
J. Leong-Skornickova
Anew lectotypiflcation of Zingiber gracile var. elatius (Zingiberaceae) 145
J. Leong-Skornickova, A. Thame & P.T. Chew
Notes on Singapore native Zingiberales I: A new species of Zingiber
and notes on the identities of two further Zingiber taxa 153
S. Lindsay & C.W. Chen
Three new combinations in Haplopteris (Pteridaceae subfam. Vittarioideae) 169
R.O. Makinson & B.J. Conn
Puccinia psidii (Pucciniaceae - Eucalyptus Rust, Guava Rust, Myrtle Rust)
- a threat to biodiversity in the Indo-Pacific region 173
D.J. Middleton, J. Leong-Skornickova & Q.B. Nguyen
Anew species of Billolivia (Gesneriaceae) from Vietnam 189
M. Moller, W.-H. Chen, Y.-M. Shui, H. Atkins & D.J. Middleton
A new genus of Gesneriaceae in China and the transfer of Briggsia species
to other genera 195
J.D. Mood, J.F. Veldkamp & L.M. Prince
A new species and a new record of Boesenbergia (Zingiberaceae)
for Thailand 207
J.D. Mood, J.F. Veldkamp, S. Dey & L.M. Prince
Nomenclatural changes in Zingiberaceae: Caulokaempferia is a superfluous
name fox Monolophus md Jirawongsea is reduced to Boesenbergia 215
L. Wijedasa, Z.Q. Shee & E. Chia
Conservation status and lectotypfication of Alangium ridleyi (Comaceae)
in Singapore 233
BOOK REVIEWS
Flora of Peninsular Malaysia. Series II: Seed Plants, Volume 4. R. Kiew,
R.C.K. Chung, L.G. Saw & E. Soepadmo (eds). 2013. (Y.W. Low) 241
A Guide to the Native Palms of Singapore. (Adrian H.B. Loo, Ang Wee Foong,
William J. Baker & Hugh T.W Tan). 2014. (K.M. Wong)
243