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Willdenowia 40 – 2010 205

Siti Nurfazilah bt Abdul Rahman1, Ahmad Sofiman Othman1, Mohd Fahmi bin Abu
Bakar2 & Peter C. Boyce1*

Studies on Hanguana (Commelinales, Hanguanaceae) for Sunda II: Five new forest
species from Peninsular Malaysia and recircumscription of Hanguana malayana

Abstract
Siti Nurfazilah A. R., Ahmad Sofiman O., Mohd Fahmi A. B. & Boyce P. C.: Studies on Hanguana (Commelinales,
Hanguanaceae) for Sunda II: Five new forest species from Peninsular Malaysia and recircumscription of Hanguana
malayana. – Willdenowia 40: 205 –219. – Online ISSN 1868-6397; © 2010 BGBM Berlin-Dahlem.
doi:10.3372/wi.40.40206 (available via http://dx.doi.org/)
Field work in Peninsular Malaysia as part of the first author’s research into relationships and evolutionary polarity be-
tween Hanguana malayana, a widespread freshwater helophyte, and the numerous and mostly undescribed Sundaic
forest species has to date resulted in the collection of five distinctive novel forest-dwelling Hanguana species. These
are described here and figured in colour. Increased understanding of these and other forest species of Hanguana has
enabled a concise delimitation of the taxonomically long-obfuscated colonial helophytic H. malayana, and this is
here presented, and the species figured. The current taxonomy of Hanguana is summarised, and a key to the so-far
described Peninsular Malaysian species is provided.
Additional key words: monocots, taxonomy, mesophytes, helophytes

Introduction
Although the taxonomy of Hanguana (Hanguanaceae) Hanguana Blume, Enum. Pl. Javae: 15. 1827.
in Sunda remains beset with considerable confusion, Type: Hanguana kassintu Blume
field work in Malaysia is beginning to permit better un- =  Susum Blume ex Roem. & Schult., Syst. Veg. 7(2):
derstanding of the many taxonomic novelties and through 95. 1830. – Type: Susum anthelminthicum Blume ex
this gradually establish a stable taxonomic platform from Roem. & Schult.
which to begin phylogenetic investigations of the genus =  Veratronia Miq., Fl. Ned. Ind. 3: 553. 1859. – Type:
Hanguana (Nurfazilah & al., in press). Veratronia malayana (Jack) Miq.
Aside from making a start to bring taxonomic clar-
References. — Backer 1924: 3, 1951: 248 – 250; Airy Shaw
ity to forest species of Hanguana, ongoing field work in
1965: 260 – 261; Rudall & al. 1999: 311 – 330; Tillich &
Peninsular Malaysia has facilitated correction of several
Sill 1999: 215 – 238; Dassanayake 2000: 214 – 215.
morphological misconceptions, ecological inexactitudes,
and taxonomic and nomenclatural problems associated
1. Hanguana bakoensis Siti Nurfazilah, Sofiman Othman
with Hanguana malayana (Jack) Merr.
& P. C. Boyce, in press. – Holotype: Malaysia, Sarawak,
Bahagian Kuching, Bako N. P., Lintang Trail, 27.5.2007,
Results Nadiah I., Malcom D., Army K. & al. S.100599 (SAR!;
Taxonomy of Hanguana isotype: KEP, n.v.)

The changes and novel additions proposed in this paper 2. Hanguana bogneri H.-J. Tillich & E. Sill in Sendt-
enable the current taxonomy of Hanguana, comprising nera 6: 216. 1999. – Holotype: Malaysia, Sarawak, bei
10 species, to be summarised as follows: der Orang-Utan-Station Semenggoh, J. Bogner 94/2211;

1 Pusat Pengajian Sains Kajihayat [School of Biological Sciences], Universiti Sains Malaysia (USM), 11800 Pulau Pinang, Ma-
laysia; *e-mail: phymatarum@gmail.com; hp:/mobile/cell +6 017 329 2609 (author for correspondence).
2 Department of Biotechnology, Faculty of Agriculture and Biotechnology, Universiti Darul Iman Malaysia (UDM), Malaysia
Kampus Kota, Jalan Sultan Mahmud, 20400 Kuala Terengganu, Malaysia.

stigma lobes flat. .) Masam. (L!. In particular he not- is obvious from the protologue discussion (Blume 1827) ed the marked disparity in size between the inner larger and from examination of the type collection that H. . Infructescence with caducous foliaceous bracts. tial inflorescences each with 2 or rarely 3 branches. with the whole 3 – 4  mm diam. Bekn. . Mesi- Key to peninsular Malaysian Hanguana species lau River. . ..10. ma­ some detail. . – Holotype: Malaysia. . spreading in fruit. Hanguana kassintu Blume. 1500  m [converted from feet on the label]. . malayana inadvertently included in the original publication awaits – Leaves arching. Veg. . also the much smaller fruit (“baccis multo minoribus”). . . – Holotype: In- Susum minus Miq. in Philipp. 7(2): 1493. 1915 ≡ Veratrum malayanum Jack. . present paper. (M!). 1830 . .   H. Blume s.2. 3. . Susum malayanum f. . Fl. sylvatica Backer. . . Leaves stiffly erect. .206 Nurfazilah & al. pro parte. are familiar.: 598. .   2 Notes. Pl. . . . isotype: BO!). 4. . Boyce. plants de- Note. Sofi. Java 3: 3. pantiensis =  Hanguana aquatica Kaneh. . who. . . malayana. fruits globose with a briefly stipi- . In fact. present paper. pointed. under the synonymy of H. the sepals to be “laciniae” and sintu is a forest species fully distinct from H. . . 35: 819. man Othman & P. . kassintu auctt. Boyce. . . . donesia. 1935.. connate at base. Syst. . a paper deal. man (‘prov. it is a feature (1827)”. Sofiman Othman & P. In the contin- That H. Chew & Corner RSNB 4233 (K!). . nitens C. As yet end of the fruit. . kassintu (Blume) Insertae sedis Backer. and almost obscuring the one (Chai S 34089) not cited in the material seen. Ind. . branches of the partial inflores- cences spreading and forming regular tiers. 5. H. . . . . .. ripen- completion of this examination. vix Bl. . . . . 6. ing with the taxonomy of H. . . . Javae: 15. Hist. . . . but provided no detailed discussion. Although assuming Miquel’s “H. Nat. resents two additional and furthermore novel taxa. . long-attenuate. . . . Ned. Fl. separate. . . 5. . fruits ellipsoid. 1820 – Stigma terminal  . . – Fertile portion of infructescence far exceeding leaves. . — Backer (1951) followed Merrill (1915) in Notes. 1. not yet traced). Stigma inserted obliquely  . . Eerste Bijv. . Boyce. . stigma lobes small. it minthicum (= Hanguana malayana). margins of freshwater bodies. . C. . & Schult. . Schult. . Hanguana exultans Siti Nurfazilah. Hanguana major Airy Shaw in Kew Bull. . 1924 ≡ Hanguana malayana subsp. . . . . cited: ascribe Miquel’s epithet. . – Holotype: Indonesia. . in Trans. Sumatera Tengah. . leafless stem up to 1. Hanguana nitens Siti Nurfazilah. 9.: 81. . . 1949.   3 Hanguana major. . provided in the present paper below. . . . Priaman’) (BO?. . . . . glossy purple-red. The plant is also noted as being glabrous. . . Mohd Fahmi. .   5 =  Hanguana anthelminthica (Blume ex Roem.. . . . Boyce. .1996 8. . L?. . 1827 10. . . 1(1): 25. . . Born. . . . . . plants sessile even in old age  . .: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia kultiviert im Botanischen Garten München. . . 1981. . . . Fl. Sabah. . Plants of shaded peatswamp mires . C. present paper. . . . . . . Plants of open situations along muddy we have not examined all the cited material. Pl. present paper. major and the other species and of freshwater swamp forest  . Hanguana podzolicola Siti Nurfazilah. ripening tion does not feature any type elements and in fact rep. Javae: 15 observation of laciniate tepals is correct. . Hanguana stenopoda Siti Nurfazilah. . . Mohd Fahmi. 21. . Mohd Fahmi. was perfectly aware of this. . . Sofiman Othman & P. & 4. . par- 7. . 10: 3. kas­ and outer smaller sepals. 2. ascending in fruit. acute. We are therefore in no doubt that it repre- sents a distinct. Phan. Blume. Soc. . banks of large rivers. .. Sofiman Othman & P. . J. who knew both taxa. Fertile portion of infructescence not exceeding leaves. ing semiglossy black.   H. C. almost certainly forest-dwelling taxon. kassintu and the other Javan species will be otherwise not recorded in any Hanguana with which we the subject of another paper in this series. The plate accompanying the descrip.   4 Misc. . . . . Enum. Enum. Boyce. . . .n. . erect. . Mohd Fahmi.5 m tall  . Java “crescit in sylvis montium Seribu Javae in- 1861. comparing it against Blume’s Susum anthel­ layana. . branches of the partial inflorescences sum anthelminthicum Blume ex Roem. malayana. . . Stoloniferous colonial helophytes  .1964. Paria- sulae”. . Handb. . . Mohd Fahmi. Enum. . Java 10(212): 2. Malayan 3. Kinabalu. . . podzolicola 5. . — A full synonymy and revised circumscription is veloping an erect. . C. .. — Miquel’s protologue describes this plant in assigning Hanguana kassintu as a synonym of H. Notes. ≡ Susum kassintu (Blume) Kurz in Flora 56: 224. . . . fruits globose. Sci. Hanguana malayana (Jack) Merr. . . . . 1873 ≡ Sofiman Othman & P.   H. C.   H. . . . . . . . present paper. . . — There are problems with the circumscription of – Clumping mesophytes lacking stolons  . . kassinutu is distinct is also implied by Airy Shaw ued absence of an authentic specimen it is impossible to (1978). Hanguana pantiensis Siti Nurfazilah. panicle very open. . . Formosa 25: 8. 1942 ≡ Su­ panicle dense.

. in rather fanciful allusion to this species remaining aloof the margins somewhat sharp. both 1 – 1. . thyrsoid-paniculate. thyr. sp. not exceeding the Hanguanae malayanae superficialiter habitu stolonifero leaves. – Fig. 19. conspicuously raised. petiole very pronounced. C. these often fewer. lamina conspicue longa scape together up to 60 cm tall.Willdenowia 40 – 2010 207 tate stigma. pseu­ dopetiole up to 80 cm long. lobes connate basally. ceous bracts. tips arching. moist mixed dipte. 25 cm) infructescences similar to that marking the start of the slender. tial infructescences each comprising up to 7 branches.2''E. .   H. stigmatis lobis separatis erectis acutis to 9  cm long. ventrally gibbose-ellip- ventraliter gibboso-ellipsoideis. Johor Bahru. 103°57'20. .2010. vainglorious. from the more abundant and parapatric Hanguana pan­ wards. Hutan Li- pur Panti. Boyce & Ooi Im under closed-canopy lowland humid moist peatforest. 40 m. somewhat flocculose abaxi. apice longe attenuato. deep chocolate sterile.5 cm. apex long-attenuate. Plants of well-drained slopes scape.9 mm. somewhat ange brown lobes.9 × 1. In­ fructescence erect. stigma sessile when fresh. Mohd Fahmi.1 mm. moist mixed dipterocarp forest rather attenuate. . petiolar tiensis. Female flowers scattered. ally.   H. — Hanguana exultans is most similar to H. silver-green adaxially. Female and male inflorescences not observed.7''E. especially the scape. lateral bose-ellipsoid. P. dry- ing deep brown. Sofiman raised abaxially. Johor Bahru. sessile. 2/3 of Othman & P. conspicuously very pale (ad 2/3 – 3/4 folii longitudine) pseudopetiolata leniter plica- brown floccose. . — Latin “exultans” = boastful. dull olive-green abaxially. tip ridges in lowland humid. . midrib somewhat prominently round. fructibus rather sharply ascending in fruit. 2. . visible portion of peduncle up seminitide nigris. partial inflorescences each with 4 – 5 branches infrutescentiis bracteis persistentibus foliaceis. . comprising 3 free man Othman & P. orange-brown lobes.5 mm wide. the largest c. sheath with margins wide. subtending bract of partial speckles. drying dark brown. dioecious mesophyte the slender. drying Hanguana exultans Siti Nurfazilah.5 × 2 cm. . cence. . 18.7''N. Notes. diminishing in size distally along the infructes- and low ridges in lowland humid. c. 55  m. connate) orange brown lobes and the fruit ripening pale aratis (non connatis) distinguenda. median branches c. Boyce based on fruiting material and observations of inflores. Mohd Fahmi. 1. whereas H. P. yellow without conspicuous black speckles. Older plants of Hanguana exultans do not develop Solitary medium-sized. There are also ecological differences: H. inner tepals c. poda although readily distinguished by the nearly trans- centiis partialibus omnibus ramis 4 vel 5. stenopoda all infructescence bracts drying pale straw-coloured. facile inflores. comprising 3 free or. distinguenda. Hin HA-55 (KEP!). ste­no­ Hanguanae stenopodae maxime similis.4. C. stigmate sessili. short. . rocarp forest on yellow clay soils  .4. the smallest 15 × 3 mm. leafless stem. . mainly solitary. drying shallowly channelled for c. 24 cm long. 1 cm wide. Boyce. broadly lanceolate. Mersing. outermost c. Hutan prominent minutely tessellate-striate in most specimens. dark green when ta. stenopoda. fruits ventrally gib. tepalis interioribus fere diaphanis. nov. comprising up to 6 partial alternate-secund. C. all tepals clasp- New forest species from Peninsular Malaysia ing the base of the fruit. sed facile foliis arcuatis (non rigide soid infructescences plus a terminal spike. base brown. leaf blade up to steno­poda is restricted to well-drained slopes and low 90 × 15  cm. soon glabrescent. drying on yellow clay soils. Ripe fruits ventrally gibbose- ellipsoid. the blade. glossy erose-marcescent. stigma sessile. 9 cm long. glossy deep green when fresh. 1. Hanguana nitens Siti Nurfazilah. arising simultaneously from the axil of the rather sharply ascending in fruit. Leaves up to 1. fruit ripening pale yellow without erect. Simpanan Lenggor. . — Low-lying wet (but not swampy) podzols Siti Nurfazilah bt Abdul Rahman. almost certainly erect at anthesis. although. to c. & Ooi Im Hin HA-48 (KEP!). bases imbricate. shallowly channelled.2 mm diam. branches up to 5 cm long. fruit ripening white with conspicuous black clawed. perianth of 6 trans- podzols in peatforest  . infructescence with persistent folia- valde ascendentibus. C. Boyce.8 m long. almost 1/2 the entire leaf length. sp. . 1. . naked stem typical for H. semi-erect. 52 × 10. longitudinally folded in. peduncle and erectis) graviter nitide viridibus. plant of low-lying wet podzols in peatforest. ovate. pale yellow. bract marking start of scape foliaceous. up to 15 togeth. accounting for Etymology. Holotypus: Malaysia. herbaceous. subtending bract.7''N. persistent. Kota Tinggi. nov. very pale green tepals. exultans parent.. 20. flush distally adaxially and all other venation Holotypus: Malaysia. par­ – Infructescence with persistent foliaceous bracts. 2°15'72. coloniali similis. Siti Nurfazilah bt Abdul Rahman.1 × 1.2010. 1°48'07. exultans is a er. 103°43'76. lobis 3 sep. Plants of low-lying wet with an associated minute bracteole. Sofi. his fructu satis parent inner tepals. cence architecture. partial inflorescences each with 4 – 5 branches. Seeds not observed. all conspicuous black speckles. old plants developing a short (to c. base long-decurrent. maturatione dilute luteis soid fruits with a sessile stigma comprising 3 separate (not sine punctis conspicuis nigris. 1 m tall. c. fructibus globosis maturatione fresh. elongate-elliptic. ovate. Ecology.

bract subtending & Ooi Im Hin HA-51 (KEP). inflorescentiae narrowly triangular. up c. dioecious stoloniferous helo. towards the tips of branches. consists of a much smaller single population olar sheath margins drying papery. the entire leaf length.2''E. 5 – 24 cm long. angle further up. peti.2010. 1 × 2. accounting for c.5 mm.35  m long. densely threatened by an adjacent building development. aetate sine cul- scape large.5 cm. 3 mm wide. P. these deep green and foliaceous towards habit is superficially similar to H. Seeds olive-green abaxially. — Blackwater mires over saturated deep peat woody. pale green with areas of Siti Nurfazilah bte Abdul Rahman. — Latin “nitens” = shining. matte black. tip long-attenuate with a conspicuous apicule Infructescence erect. Hutan diam. with age entirely glabrous. very much depleted unsexed population critically irregularly but prominently plicate when fresh.208 Nurfazilah & al. especially bus (non effusis) differt. pale green. Female flowers ± distant. and globose fruit sometimes very sparsely flocculose abaxially along the ripening semiglossy black with separate.4. The second known locality. base clasping. median branches longer than lateral branches. Siti Nurfazilah bt Abdul Rahman. fertiore. enveloped by appressed Notes. 3 mm diam.5''E. sp. P. at Hutan Lipur Panti. to 5 mm. lateral branches ap- proximately 3. 2 mm cate. brown with remaining hairs pale brown.  1. tip long-attenuate. phyte to c. Ecology. soon glabrescent. leaf blade up to 84 × 13. narrowly lanceolate. 9 cm long. C. malayana. 20. staminodes not observed. gla. Mohd Fahmi. Kota Tinggi. sion to the remarkably lustrous leaf blade. 1°48'07. ole up to 15 cm long. 4 mm diam. stigma sessile. 1/5 of curved.5 mm. silver-green adaxially. ± V-shaped in cross-section. Johor Bahru. C. – Fig. flush to slightly impressed adaxially. erect at anthesis. — Malaysia: Johor Bahru. ovary spreading. shallowly channelled. brescent or nearly so in part. P. So far. foliaceous.  1  m tall. Siti Nurfazilah bt Abdul Rahman. rectangular in cross Ooi Im Hin HA-57 (KEP). Boyce ceous. in allu- moderately stout pedunculate panicle. Hanguana nitens is known from three sites.5  cm diam. stolons up to 45  cm long (although peatswamp forest at altitudes of 40 – 50 m. moderately extensive female stands. pointed midrib. visible portion of peduncle up to 30  cm long.4. 103°54'75. folia. in. herbaceous. dark shining green when fresh. paniculae structura con- Male flowers not observed. longitudinally veined. Boyce & Ooi Im purple-brown speckles and spots when fresh.. Hutan Simpan Panti. — Malaysia. the older portions clothed in dense fibrous de. bract subtending proximal Tinggi. semi-erect or creeping. occurring as several the base decurrent on the pseudopetiole.2010. nov. narrowly elliptic. apiculate. 3-lobed. ner tepals c. outer 3 tepals c. with the blade conspicuously long- decaying into fibres. 2/3 – 3/4 of the leaf male population.. Solitary robust. C. 1. c. So­ branches weakly ascending. distal-most partial inflorescences small to minute. usually less). Kota Tinggi. sessile with a broad base in the axil of a short. broad bract. green. acute-pointed. the whole inflorescence weakly flocculose. deep chestnut brown and eventually partially or completely glossy green leaves. pseudopetiole accounting for c. sterile. ripening through arching with the tips touching the ground. with up to 15 pa­tently branched thyrsoid or spicate partial inflores. Lipur Panti. innovations weakly plicate with a long-attenuate tip. At the type locality it is abundant. 19. ± erect.. terminal spike c. 103°57'20. 9 cm wide. rhizomatous with the terminal portion ascending. Hutan Sim- length.. pan Panti. Boyce & up to 15 cm long and c. 2. Johor Bahru. Other material seen.4. margins sharp. dioecious mesophyte to nate at the base. midrib acutely raised of female plants. stigma remains matte black. tip long-atten. long-pseudopetiolate. drying deep Hin HA-56 (KEP!). base diam. decurrent. bases imbri- yellow-green to black. and a single isolated uate. all Hanguana pantiensis Siti Nurfazilah. 1 cm wide. leaf stigma lobes. pseudopeti­ hemispherical. drying dull semiglossy black. c. c. the outermost leaves broadly ovoid-globose. c. deeply excavated. parte fertili foliis non excedenti. 103°57'20.2''. 0. drying dull olive-green. occasionally in small clusters. colonial petiolar sheaths.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia Medium-sized.7''N. flocculose abaxially.. 60 cm long. 1°48'07. lobes separate. c. subtended by a fully developed foliage leaf. partial inflorescences very narrowly triangular. c.6 m long. C.5 cm diam. Leaves up to 1. up to pseudopetiolate (up to 2/3 – 3/4 of the leaf length) and 16 together. stem terete. Hutan Lipur Panti. is a abaxially. polished. glabrous. erect. Leaves up to 1. lower part of peduncle up to 1. with the pit margins in. ovate. along margins of closed-canopy lowland humid moist graded leaf bases.6''N. blade relict. 2 m tall. — Hanguana nitens by its stoloniferous. 2010.5 – 2 cm wide. the margins . erect.5  cm Holotypus: Malaysia. Distribution. 2 × 2. or burrowing through liquid peat. section at the base. but may be the stolon active tip. bract marking onset of Hanguanae podzolicola maxime similis. mo longo efoliato.5 cm long. leathery. pedunculo breviore. blade 45 – 100 × 5 – 8  cm. sub. Boyce. peduncle and scape together fiman Othman & P. older leaf portions turning glossy readily distinguished by arching (not stiffly erect). The third site. with numerous close-set very fine cross-veinlets. up to 80  cm tall. Johor Bharu. deep green semiglossy when fresh. Kota cences plus a terminal spike. Female and male inflorescences a Etymology. 40 m. tepals shortly con. 19. up to 16 together.7''. Ripe fruit globose. inflorescentiae partialis ramis fructu ascendenti- mostly solitary. 1°52'22.

Notes. pseudopetiole c. — Hanguana pantiensis is one of a group of spe. although. referring to the type and only known lo- cent. all with an associated common. At the time of collection in length. perianth of ing for up to 1/3 of the entire leaf length. ginating from. sterile. ing start of scape foliaceous. not exceeding the & Ooi Im Hin HA-50 (KEP!). nov. P. ripening glossy pink. ously very pale brown flocculose. – Fig. bract subtending partial peduncle. 2 m tall. drying dark brown. the largest c. dark brown. soon falling (but similar to H. the small. base decurrent. stem with age. 103°43'76. visible por. soon falling (but more investigation needed).  1  mm.. 55 m. lanceolate-elliptic. Ripe fruit pink. tary. minute raised yellow speckles. dense-paniculate.7''N. the leaf blade to form the apicule. ally. longitudinally folded inwards. prominent minutely tessellate-striate in most specimens. 10 – 18  cm long. leafless partial infructescences each comprising of 9 branches. green when cies. mainly solitary. Simpanan Lenggor. tending bract. 5 mm midrib pronouncedly round-raised abaxially. sterile. with age becoming leafless to 1. based on fruiting material and observations of inflorescence architecture. herbaceous.2010. perianth of 6 opaque. with the fertile portion of the inflorescence not infructescences similar to that marking the start of the exceeding the leaves. the largest c. all te. shallowly chan- 6 opaque. midrib prominently round-raised abaxially. Female flowers scattered. drying somewhat equal in length. bract seemingly mark. softer and dull olive-green abaxially. conspicu. H. lanceolate-elliptic. oc- with spent inflorescences suggested that male plants were casionally in groups of 2. stem erect. raised. diminishing in size distally along the infructes- the branches of the partial inflorescence ascending (not cence. the denser panicle structure. the smallest 25 × 5 mm. in which early in development the ovary fresh. thyrsoid or more rarely spicate infructescences plus a terminal spike. drying dark brown. praeterea individuis vetioribus caule tion of peduncle up to 20 cm long. cence architecture. subtending Solitary robust. 2 × 2. lower. erect. sessile. ovate. drying flush adaxially and all other venation Hanguana podzolicola Siti Nurfazilah. c. conspicuously very pale brown flocculose. 45 cm long.4. erecto efoliato evoluerentibus distinguenda. inner tepals c. partial infructescences each comprising up to 80 × 13 cm. Sofiman Othman & P. florescences not observed. Boyce fructescence erect. the infructescence. — Well-drained flat and slightly undulating almost certainly erect at anthesis. espe.Willdenowia 40 – 2010 209 somewhat sharp. channelled diam. somewhat several female plants were fruiting. podzolicola. Female and male in­ where it is abundant. dioecious mesophyte to bract of partial infructescences similar to that marking c.7''E. dry- connate. — Type locality “Panti” + Latin “ensis” = ori­ most part of petiolar sheath with margins erose-marces. stigma 3-lobed. spreading. pantiensis is most foliaceous. Hanguanae pantiensi maxime similis in stigmatis inser- peduncle and scape together up to 70 cm tall. spreading) in fruit and by not developing a tall. Johor Bahru. Mohd Fahmi.  1. lesser venation very obscure when fresh. — Only known from the type locality. with fresh. Boyce. Infructescence soli- areas of lowland closed canopy humid moist peatforest. and numerous plants flattened. diminishing in size distally along with a terminal crown of leaves. tip up to 12 branches arising simultaneously from the axil of long-attenuate with a conspicuous apicule to 7 mm. In­ 18. differing by the much shorter more investigation needed). drying medium-brown and ultimately marcescent. ascendentibus). nelled. up to 14 cm long. ses. ovate. minute bracteole. ing closely and conspicuously longitudinally-raised stri- served.5 × 2 mm. with the marginal veins drying darker adaxially and overtopping Distribution. Female flowers scattered. whorled. 1 – 2 mm wide. Leaves up to 1. 2°15'72. Hutan based on fruiting material and observations of inflores. comprising up to 11 partial. c. up to 15 together. 41 × 5. forming dense pure stands. Etymology. lobes not adaxially. cially in the lower middle part of the leaf blade. green tepals. all with an associated minute bracteole.  48 × 10  cm. Siti Nurfazilah bt Abdul Rahman. 2 cm wide. sp. sed paniculae structura plurime aperta. comprising up to 13 partial. especially the scape. although. bract seemingly marking start of scape obliquely to sublaterally inserted. impressed adaxially. median and lateral branches almost equal ity.5 m long. oblique-globose.5 mm. mainly solitary. along the length. 3. out- . soid or more rarely spicate infructescences plus a termi- nal spike. especially about half way weakly concave. Ecology.5 m the start of the scape. Female and male inflorescences not observed.5 mm. account- sile. bases imbricate. almost certainly erect at anthesis. peduncle and scape together up to 80 cm tall. silver-green adaxi- edly erect. Seeds not ob. all novel. petiolar sheath margins hyaline when pals clasping the base of the fruit. Holotypus: Malaysia. leaves. stiffly the subtending bract. inflorescentiae partialis fructu ramis effusis (non valde dark green when fresh. C. narrowly elliptic. with scape. 1 – 2  mm wide. green tepals. outermost c. whorled. C. median and lateral branches almost chartaceous and deep green when fresh. branches arising simultaneously from the axil of the sub- Hanguana pantiensis is abundant at the type local. ate with pronouncedly reticulate lesser venation. leaf blade est 18 × 3  mm. Mersing. thyr­ 40 – 60 m. cality. mark. visible portion of peduncle up bends longitudinally to produce a fruit with the stigma to 30  cm long. tione obliqua. the margins very sharp.

up to 25 × 6. older (outermost) leaves spreading. outer tepals Hanguana stenopoda Siti Nurfazilah. Boyce branch occasionally solitary. exultans. very oc- & Ooi Im Hin HA-53 (KEP). although. lowermost flowers of each 19. dweller) + “podzol”. the terminal one a single spike. — Latin “cola” (derived from “incola”. angled.210 Nurfazilah & al. Holotypus: Malaysia. dark brown-red.  0. 4. forest on yellow clay soils with a moderate leaf litter layer at altitudes of 60 – 125 m.5 mm. Solitary. Siti Nurfazilah bt Abdul Rahman. Female flowers mainly in Tinggi.5 m tall. sp. but may be tic. Maturitate caule brevi gracili efoliato. in fruit. stenopoda develops a short. each comprising of 3 branches. somewhat semiterete.7''N. branches arising simultaneously from the axil of Etymology. 2. 5 × 4. almost certainly erect at anthesis. Hanguanae exul tanti similis. scattered groups of 2 to 3. the cence with caducous foliaceous bracts. infructescentiis bracteis caducis fo. oblique-glo.9 mm. somewhat impressed adaxially until the bose. weakly ferring to the restricted ecology of this species. lobes connate to form a clover-leaf.. with age. — Malaysia. up to partial inflorescence branches spreading (not sharply as. lobes connate basally. subtending partial infructescences linear-triangular. bract Notes. 1. leafless stem up to 1. base clawed. tip. all clasping the base of the fruit.  1  cm wide. channelled. peduncle and scape together up in seasonally swampy peatforest. under 1/2 of the entire leaf length. globose fruits with a briefly stipitate stigma. thereafter flush.2''N. Ripe abaxially. — Lowland humid. perianth of 6 tepals. greyish flocculose. nov. although branches subequal in distal- Other material seen. to 62 cm tall. In age. raised c. lateral branches. differing by the opaque green inner tepals. c. H.3''E. 5 mm diam. marking start of scape foliaceous. partial inflores- longest leaves arching with the tips touching the ground. – Fig. ripening glossy medium pink to ma. diminishing in size distally along the infruct- cending) in fruit. — Malaysia. Seeds not observed. median branch usually longer than inhabitant. c. Johor Bahru. Boyce HA-60 (KEP!). up to 15 together. ing material. leafless stem. his fructu effusis.5 mm wide. ovate. Infruct­ escence erect. Bakar & P. pseudopet­ brown.  16  cm long. Ripe fruit compressed-globose. dioecious mesophyte to c. visible portion of peduncle up to 22. matte black. c. bright green when fresh. Pahang. all mainly sessile. Sofi­man Othman & P.5 cm long. fructibus maturitate albis punctis conspicue nigris. mid-point. Hutan Lipur Panti. a raised peat deposit.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia ermost c. all clasping fruits in fresh material and medium green. when fresh somewhat slender. with the margins very sharp.2010. 40 – 60 m. 5 mm wide. matte deep brown. cences each with 2 or rarely 3 branches. developing a short (up to 25  cm). ob- genta. — Malaysia. partial infructescences erect. tip attenuate. with all raised. and by older individuals developing an escence. C. 103°19'83. based on fruit- Distribution. c. Leaves up to 1 m long. 7 × 2 mm. C.8 mm. moist mixed dip­ c. Boyce. leaf blade up to 51. Female and male inflorescences not observed. stigmate breviter stipitato lobis basaliter connatis graviter Ecology. lateral branches approximately 2/3 of the length of median branch. known only from the bus opace viridibus. herbaceous. ovate. drying somewhat prominent.2 × 1. erose-marcescent terocarp forest on yellow clay soils with a moderate leaf leaf tip long-attenuate. narrowly ellip- pantiensis by the oblique insertion of stigma. Pahang. 1. Taman Negeri outer with a narrow dark red margin. c. stigma 3-lobed. thyr­soid partial in- Ecology.2 mm. sterile.5  cm. – Fig. Re. the the subtending bract. mid­ litter layer. type locality. 1°48'07. c. drying strongly Hanguana stenopoda is restricted to well-drained longitudinally folded. very shortly stipitate.4. occasionally on raised podzols developed foliage leaf. especially in the lower part of the leaf blade. Kota most partial infructescence. 2 × 2 mm. P.2''E.5.5 × 2.9 mm diam. glossy Siti Nurfazilah bt Abdul Rahman. 5 – 7 cm long. all with an associ- ated minute bracteole. elliptic. inner only red at the Endau Rompin. — Hanguana stenopoda resembles H. inner tepals c. and the fruit ripening white with conspicuous iole 35–44  cm long. scure when fresh. ovate. margins rather wide. ovate. inflorescentiis partialibus omnibus ramis 2 raro 3. moist mixed dipterocarp badiis differt. in thinly leathery. 103°57'20.5  mm. casionally pedicellate to c.2010. accounting for just black speckles. Distribution. Johor Bahru. drying which the lobes are connate basally and deep chocolate thinly chartaceous and pale straw-coloured. 5. distally flush. Mohd Fahmi Bin Abu white with conspicuous black speckles. lowermost part of petiolar sheath slopes and low ridges in lowland humid. 2°37'37. ini. other venation striate. slender leafless stem Notes. bract distinguished by the much more open panicle structure. minor venation minutely tessellate.. glossy. Seeds not observed. these spreading bases imbricate. subtended by a fully humid moist peatforest. 1. — Hanguana podzolicola closely resembles H. Rompin. 1. infructes- tially semi-erect. 1 m tall. — Raised podzols in closed canopy lowland fructescences plus a terminal spike. weakly concave. 65 × 6 mm.3 mm. comprising 4 partial. stigma 3-lobed.5 × 9  cm. 12. . 1 × 1. Mohd Fahmi. C. the smallest c. tepalis interiori. liaceis. tipped with a 1 cm tubule. inner tepals rib prominent proximally. fruit pale yellow with minute pink speckles.

medium straw-coloured. lower the infructescence. 13. branches initially ascending. 1942 ≡ Su­ lateral branches approximately 29  cm long. each subtended by a semipersistent bract up to 20  cm pedunculate and scapose bracts sessile or subsessile with long. base clasping. 1935. ex Hook. enveloped by and inner tepals c. & florescences 30. with longitudinal slits.1  mm sum anthelminthicum Blume ex Roem. 1915 ≡ Veratrum malayanum Jack. peduncle up to 3 cm diam.. passing into fertile bracts. soon glabrescent or nearly so. Temerloh. fertile.5 – 2 m. with numerous close-set thin cross- Hanguana malayana recircumscribed veinlets. c. partial male in­ =  Hanguana anthelminthica (Blume ex Roem. comprising up to 10 partial infructescences flowering plants) remote. Pulau Pinang [“Poeloe Pinang”].. – Holotype: Caroline Islands.5 mm appressed sheath. 4 mm wide.  2. drying spongy-chartaceous and Other material seen. shortly less). lateral branches approximately 11  cm ex Roem.) Masam. Hatusima bract marking onset of scape large. sterile. 7(2): 1493. Palau. stem and 3 inner tepals c. exceptionally reaching 4  m. isotype: BO!). Female flowers with outer 3 tepals c.5. leaf blade ­(20 –)45 – 200(– 300) × 4 – 15  cm. 1(1): 25. India 6: 391 1892. green. scape together often more than 2  m tall. spongy. fornicate. 3 others tionally up to 3. — Malaysia. inserted on the base of the perianth. neously from the axil of the subtending bract. branches arising simulta- type: Malaysia. Boyce HA. medium to dark green. gradually narrowing into the peti- in podos = a foot. Sci. 2 mm long mud. between the longitudinal veins very densely and finely longitudinally striate. tip acute. deeply di- flocculose abaxially when young. late. – Holo. site the outer sepals minute. aquati­ cences. Siti Nurfazilah bt Abdul c. 10: 3. weakly angled. Java 3: 3 (1924) ≡ Hanguana of flowers. robustly rhizomatous with the terminal portion as. later drooping malayana subsp. partial female inflo­ & Schult. Blume s. in Blumea 6: 399 1950. the smallest c. smaller. stem clasping. pseudopetiole. in Philipp. with up to 10 whorls of 1 – 10 × Fl. less. rounded. rather distinctly yellowish. staminodes 6. c. about as long as the inner tepals. 1949 rescences stout and short. Enum. base acute to decurrent on the Gunung Benom. 1859 ≡ Susum malayanum (Jack) patently branched thyrsoid or spicate partial inflores- Planch. narrowly triangular. 1. pseudopetiole accounting for 102°12'84. Babeldaob. ellipsoid.6''E. on terete. — Ridley 1907: 131 – 133. stolons up to 2. stamens 6. ≡ Hanguana malayana var. 1/5 of the leaf length.: 81. mar- Rahman. Flowers of both sexes ± distant. Bekn.  10 × 2  cm. Ned. Ripe fruit leaf sheaths long and broad.. Fl. which finally dissolve into fibres.  4  mm wide. branches subequal in distal-most units. broadly lanceolate-elliptic. stigmas 3.) Bakh. c. somewhat spongy-leathery. although 3717 [K-WAL!]. 2 cm diam. small.4. 1 mm. foliaceous. – Epitype (designated here): Wallich EIC branches usually longer than lateral branches. (L!. on the anterior side. tip long-acute. blade densely lon- gitudinally veined. slender stem that ole. median (not traced). inserted degraded leaf bases. although these diminishing in size distally along a broad base. otherwise glabrous. – long and c. Male flowers with 3 outer tepals c. Krau Wildlife Sanctuary 3°49'29. c. & Schult. ascending. dorsally compressed. 3: 553. c. up c. sublustrous when fresh.5  m long (usually in a basal cleft.8''N. not or only very obscurely Hanguana malayana (Jack) Merr. Java 10(212): 2. Backer either solitary or in small clusters. long.5 cm long. up to 20 together. 1 mm long phytes to c. Nat. 1820 ≡ Veratronia malayana (Jack) Miq.5 m long. Airy Shaw 1975: 1 – 50. 3 mm wide.  1 × 5  mm. 13. wide. lanceo- develops in older individuals. 125. Java “in paludibus circa Buiten.) Backer. 1924: 369. C. Brit. or floating on the surface of water. Malayan culate panicle erect at anthesis and subtended by a fully Misc.n.. 0. developed foliage leaf. Hist. deeply split dark shining purple-red. stiffly erect. S.. — Greek FJg<@H (stenos) = slender. higher leaves (of cence erect.2010. up to 20 cm long. 1830 ≡ Susum malayanum f. later marcescent. um green. peduncle and Holo­type: Indonesia. plicate. midrib rather thick. dotted red. with a greater number ca Backer. visible portion of peduncle up to 50 cm long. 3 mm long. referring to the short. the base of the perianth.n. Female and male inflorescences a stout pedun- C. and Lat. on shorter petioles. the older portions clothed in dense fibrous anthers small. these foliaceous towards the stolon ac.  2  mm long. 18  cm long. those oppo- tive tip. stigma . Almonogni. f. much longer and broader. although usually somewhat filaments filiform from a broader base. gins sharp. & Schult. green or yellowish or the inner tepal Large to massive colonial stoloniferous dioecious helo. ± flat to shallowly V-shaped. Ind. anthelminthica (Blume weakly angled. Schult. Jack s. pistillode small. occasionally fleetingly ovary broadly ovoid-globose. lowermost part of petiolar sheath margins 61 (KEP). cences plus a terminal spike. at post-anthesis prior to withering. Mohd Fahmi Bin Abu Bakar & P. Handb. Pahang. Syst. Born. the whole weakly to rather =  Hanguana aquatica Kaneh. sessile with a broad 1951: 248 – 250. broad bract and tepals shortly connate at the base. cending. vided into 3 spreading broadish short arms..1938. creeping or burrowing through li­quid clavate. Fl. c. lower part of zorg”. c. or 4866 (FU!). Phan. J. References. base in the axil of a short.. Leaves excep.  3  m tall.5 cm long.Willdenowia 40 – 2010 211 Etymology. Soc. densely flocculose. anthelminthica (Blume ex Roem. stigma sessile.. c. Fl. in Trans. medi- Formosa 25: 8. Infructes­ lower leaves briefly pseudopetiolate. thinner and longer than those of female inflores- Veg. more usually about 1. erect. 15 cm diam.

. Europe in 1824”. we have opted to epitypify Wallich EIC Morphologie und Anatomie von Hanguana Blume 3717. D. Batavia: Ruygrock & Co. R. with Jack’s herbarium. land. Stevenson D. Fla­ tralia and north to the Philippines (Luzon). W. A revised handbook to the flora of Ceylon Peninsular Malaysia was in need of critical study. N. Rudall P. (1999). 1924: Handboek voor de flora van Java 3. L. the ripe fruit is dark shining purple-red. that the Flora Malesiana account for Hanguana (Backer Blume C. 214 – 215 It is interesting to note that Ridley (1907. while for the open habitat helophyte he used the Ridley H. 1978: Three interesting plants from the western Micronesia (Palau). (in press) have highlighted Djakarta: Noordhoff-Kolff. G. However. 1915: New or noteworthy Philippine plants Susum malaynanum to the forest species (in the broad XI. 1827: Enumeratio plantarum Javae et insu- 1951) contains numerous misconceptions with the result larum 1. 1907: Materials for a Flora of the Malayan (now synonymous) S. Airy Shaw H. A. [CrossRef] Distribution. P. ae). Syst.. 1924) was in: Dassanayake M. 1999: Diligent searches of relevant herbaria have failed Structure and systematics of Hanguana. — Equatorial tropics from Sri Lanka to Airy Shaw H.nationaalherbarium. (ed. Ridley (1907) unaccountably applied the name Merrill E. Peninsula 2. van (ed. matte black. K. that the circumscription of H. 12: Hanguana malayana]. Cowan (1954) “methodically searched the Herb. Field and Jack collections in the herbarium. malayana has been subject Cowan J. It might be argued that the type is lost and notes on critical morphologies for elucidating and thus neotypification is justifiable. No record of how and when they were acquired FMcollectors/J/JackW. & Clayton W. and of freshwater swamp forest. mit der Beschreibung einer neuen Art. – Pp. G. Thanks go to J.nl/ 1 dupl.212 Nurfazilah & al. Dassanayake M. As Tillich H. tunately.htm was found …”). 18: 249 – 273. – Acta Phytotax. a Pinang collection as was Jack’s. 1999: Systematische Studien zur a compromise. Edinburgh. Fosberg F. lobes connate to form a clover-leaf almost ob. for the seventh edition of Willis’s ‘Diction- ary’. van 2006: Cyclopedia of Male- (e. J. References scuring the end of the ovary. Flora malesiana. — Nurfazilah & al. 4(3). R. a mono- to turn up Jack’s collection of Veratrum malayanum [≡ cotyledon of uncertain affinity. D. 33: 1 – 5. [in press]: Studies on Hanguana (Commelinales- part of his botanical notes and collections was lost by the Hanguanaceae) for Sunda I: Hanguana bakoensis. & Boyce P.-J. – Notes. A. not least because “an important C. M. This Ridley H. circumscribed. Bot. & Sill E. south as far as northern Aus... as defined in the above key. As alluded by Steenis-Kruseman 311 – 330. 10: 1 – 84. Malaysian Borneo.g. Balkema. nov. ser. we are reluctant to take this step.. N. sense). F. Northern Territory of Australia (Thymelaeaceae. K. K. Jack material periodically turns up in ‘expected’ places Steenis-Kruseman M. D. Backer C. . Pen- ang. – Kew Bull.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia 3-lobed. names. Gard. – Leiden: Nationaalherbarium Neder- inburgh [E] for Jack specimens and found 64 in all (incl. a fire of the ship ‘Fame’ on which Raffles embarked for new forest species from Sarawak. stated by Dassanayake & al. – Singapore: Government Printer. – Fig. Unfor. ser. published at http://www. 1. – Kew Bull. – Rotterdam: A. & Linder H. Absent from courtiaceae & Hanguanaceae). Hanguana bogneri spec. – Notes Roy.). 2000: Hanguanaceae. – Sendtnera 6: 215 – 238. Bot. given that Hanguana taxonomy. 35: 819 – 821. Sci. [CrossRef] (2006) there are difficulties and obscurities associated Siti Nurfazilah A. [CrossRef] Ecology. 14.). the area between the Wallace and Webber lines. 1924: Flora of the Malay Peninsula 4. clearly aware that the circumscription of Hanguana in D. Backer C. 1954: Some information on the Menzies to considerable and quite unnecessary confusion. anthelminthica. margins of freshwater bodies. – Pp. ma­ Garden. A. Royal Botanic observations in Malaysia leave us in no doubt that H. J. Geobot. as London: Reeve & Co. Ed. Veldkamp (Leiden) for translation of the Latin diagnoses. C. Acknowledgements Fieldwork in Malaysia was supported by a Research Uni- versity Grant from the Universiti Sains Malaysia. Edin- layana is both morphologically and ecologically clearly burgh 21: 219 – 227. – is incorrect. c. — Plants of open lowland situations along mud. 3 mm diam.. etc.). Backer (1951) stated the fruit to be shining red. – Leiden: Van Leeuwen. 1981: A new species of Hanguana from Borneo. J. – Austr. 1965: Diagnoses of new families. and beyond any (Hanguanaceae) und Flagellaria L. – Kew Bull. – Philippine J. Airy Shaw H. – dy banks of large rivers. 1951: Hanguana. sian collectors. 6 – 7. Ahmad Sofiman O. new Seeds not observed. 248 – 250 in: Steenis C. (Flagellariace­ doubt well-representative of the species.

Hanguana exultans – A: flowering plant in habitat. . – A – D: Siti Nurfazilah & al. D: median leaf and inflorescence of H. C: detail of the ventrally gibbose-ellipsoid fruits ripening pale yellow with a sessile stigma comprising 3 separate (not connate) orange brown lobes. exultans (right). note the rather sharply ascending branches. images © Rosazlina Bt Rusly. pantiensis (left) compared with H. HA-55. 1. B: infructescence.Willdenowia 40 – 2010 213 Fig.

C: plicate leaf blade. pointed stigma lobes with those of H. erect. – A – E: Siti Nurfazilah & al. long (2/3 – 1/4 of the leaf length) pseudopetiole. 7B). E: ripe fruits. note the arching (not stiffly erect) leaves. D: portion of an immature infructes- cence.: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia Fig. malayana (Fig. HA-48. compare the separate. . B: portion of a mature plant showing a stolon and the conspicuous.214 Nurfazilah & al. Hanguana nitens – A: plants in habitat. 2. images © Rosazlina Bt Rusly.

. B: detail of infructescence. D: leaf bases showing litter-trapping. HA-56. note the dense panicle with branches of the partial inflorescences ascending. note the dense panicle carried down in the leaves. with the stigma obliquely to sublaterally inserted by longitudinally bending of ovary. images © Rosazlina Bt Rusly. C: ripe fruits. Hanguana pantiensis – A: plant in habitat. 3. – A – D: Siti Nurfazilah & al.Willdenowia 40 – 2010 215 Fig.

: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia Fig. C: ripe fruits. note the open nature of the panicle and the spreading branches of the partial inflorescences. images © Rosazlina Bt Rusly. – A – D: Siti Nurfazilah & al. Hanguana podzolicola – A: plant in habitat showing the tall. D: median leaf and inflorescence. 4. HA-49. with the stigma obliquely to sublaterally inserted by longitudinally bending of ovary.216 Nurfazilah & al. leafless stem produced by older plants. B: detail of infructes- cence. .

D: ripe fruits with opaque tepals. images © Siti Nurfazilah Bt Abdul Rahman. Hanguana stenopoda – A – B: plants in habitat. . note the partial inflorescences each with only 2 or rarely 3 branches and that the subtending bracts have fallen. and the fruit with conspicuous black speckles. 5. briefly stipitate stigma. B: detail of infructescence. with the lobes connate. – A – D: Siti Nurfazilah & al.Willdenowia 40 – 2010 217 Fig. deep chocolate brown. HA-60.

P..: Studies on Hanguana for Sunda II: Five new species from Peninsular Malaysia Fig. 6. .218 Nurfazilah & al. – A: image © Mike Lo. Sarawak. note the stiffly erect leaves and the stolons floating on the water surface. B: image © Siti Nurfazilah Bt Abdul Rahman. Perak. Hanguana malayana – A: plants in habitat in Maludam N. B: detail of semi-terrestrial stolons in habitat.

more slender rather yellowish branches of the male inflorescence. note the much longer. 7. the ellipsoid fruits. and the large sessile stigma almost obscuring the end of the fruit. C – D: male inflorescences. the green partial inflorescences. robust branches.Willdenowia 40 – 2010 219 Fig. note the short. – Images © Siti Nurfazilah Bt Abdul Rahman. Hanguana malayana – A – B: female plants. . after anthesis these branches droop prior to withering.