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ISSN 1346-7565 Acta Phytetax.Geobet. S6 (]):

41-53(200S)

Return from the Lost: Rediscovery of the Presumed Extinct

Leptosolena(Zingiberaceae) in the Philippinesand itsPhylogenetic
Placement in Gingers

HIDENOBU FUNAKOSHI]*, W. JOHN KRESS2, JANA gKORNIeKOVA3,

AIZHONG LIU2 and KEN INOUE`'

iDepartment
qf'Environmental System
Science,GraduateSchoolofScience
and TlechnologM Shinshu U}iiversiijl
3-
2Dapartment
1-lAsahi, Matsumoto 390-862J Japan; ofBotany
MRC-166, UhitedStatesNdtionalHerbarium,
IVationalMuseum ofNZitural HistorplSmithsonianlnstitution,R O. Box 37012, Ukeshington,D,C 20013-7012
3Department
USA; ofBotan>LCharles University,Bendtskd 2, J28 Ol,Prague,Czech Rqp"hlic;4Biolegicat
Institute and Herbarium,fuculty ofScience,Shinsht{Universic)l3-1-1 Asahi, Matsumoto 390-8621Jopan

The genus Leptosolena currently accepted as monotypic and endemic to the Philippines, has beencon-
sidered as an imperfectly known genusdue to the description basedon insucacientherbarium materi-
als fOrdescribing fioralcharacters and no recent collection. Our rediscovery of L, haenkei has made it
possible not only to describe the species in more depth from fresh materials and to compare with the
uncertain second species, L. insignis,more precisely, butto clarify the phylogenetic position ameng
Zingiberaceae with molecular data. Our results support theformertreatmentthatL haenkeiand L insig-
nis are conspecific, resulting in L. insignisas a latersynonym. The ]ectotype of L. haenkeiischosen
among Haenke'shistorical colLections deposited at PR and PRC. Resultsfrom DNA sequence dataof
the ITS and tnatK locidemonstratethatLqptosolena forms a clade with LEiizoverberghiaand Aipinia
species from the Philippines and Oceania.

Key words: ITS, lectotypification,

Leptosotena, Leptosoienahaenkei,matK, molecular phylogeny,
Philippines,
rediscovery, Zingiberaceae

The genus LeptosotenaC. Presl (Zingiberaceae)

length (Smith
1990).The taxonomic positionof
endemic to Northern Luzon, Philippines,
compris- this curious member of Zingiberaceae is always
es only one L.haenkeiC, Preslas current-
species, controversial. Bentham (1883) and Burtt & Smith

ly accepted (Larsenet al. 1998).Leptosolena is (1972)mentioned a possible close aranity with

outstanding inZingiberaceae by thelargeflowers Burbidgea which shares very short filamentand

with extremely longand slender corolla tube which smaller labellumthan corolla lobeswith Lepto-
are exerted from the calyx for more than halftheir solena. Schumann (1904) based on Presl(1827)'s

* Author forcQrrespondence: E-mail: tO lh1 1 I @amail.shinshu-u.ac .jp

tProfessorKen inouewas ki11ed
by accident duringfield work in Sakhalin.The present
paperisdedicatedfor the inspir-
ing memory of the lateProfessor
Inoue.

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42 APG Vbl.56

and linedrawingtransferred Leptoselena

description discussabout L. auricutata not in the present study.

under the genus Aipinia L., subgenus Autaipinia Leptosolena haenkeiwas recollected from the
K, Schum. and established section Leptosolenia valley slope along Chico River near Bontoc,
(C.Presl)K. Schum. He changed sucax of Mountain Province, Northern Luzon by the first
"Lepto-

sotena" into `CLeptosotenia"

probably due to fbl- author and Leonardo Co in May 2002. The redis-
lowing the orthodox Latinword forrnationfrom covery of the plantsin fu11bloom in itsnatural

the Greek word habitat hasmade itpossible not only to describe the
"lepto-solen" "slender
meaning

pipe." Ridley(1909) recognizedLfu)tosotena as a species in more depth and to compare with an

distinct genusconsidering alliance with Hkdychium unceitain second species, L. insignisRidl,,but to

which shares longcorol]a tube with Lepto-solena. carTy out DNA-sequencing studies to clarify the

Burtt& Smith (1972) pointedout that L. haenkei i's phylogeneticrelationship of this genus with other
unlike any otherAipinia species becausethe inflo- generaof subfamily Alpinioideae of the Zingiber-

rescence lacksbractand bracteole, and the corolla aceae.

tube isextremely long and narrow, thus suggesting

to maintain Leptosolena at generic levelpending Materialsand Methods
more critical study. Lasrsen et al. (]998) mentioned

poorlyknown genus,endemic to The field collections and tissue

"A
this genusas samples were made

the Philippines, Luzon, with L. haenkei as the only in the Mountain Provinceof Northern Luzon.
species." Kress et al. (2002) proposeda new supra- Herbarium material examined was from the fo1-
generic classification of Zingiber-aceae based on lowinginstitutions: AAU, BR, E, GH, KYO, L, P, PNH,

molecular data,although Lepto-solena, good tis- PR, PRC, PUH, SING, and us.
sue materials beingnot available, was tentatively Data from previous investigations of the phy-

placedunder subfamily Alpinioideae Link, tribe logenetic relationships within the familyZingiber-

Aipinieae A. Rich. based on morphological fea- aceae (Kress et al, 2002)were used to determine the
tures. evolutionary positionof the genus Lqptosotena in
In 19C}6, Ridley describedthe second species of the subfamily Alpinioideae.In total 5 1 gpecies were
the genusLeptosolenainsignis Ridl.based on A. D. analyzed, includingthree outgroup taxa inthegenus
E. Elmer'sspecimens collected at 1[iwin Peaks (in Siphonochitus, 46 taxa previously sequenced in the
Municip. Tuba near Baguio City),Benguet Proyince Alpinioideae,and thetwo new taxa. Comparative
in Northern Luzon (Ridley1906). But Merrill sequence data of the internal transcribed spacer

(1925)
rendered the species synonym of L. haenkei (ITS)lociand matK-trnK fiankingintergenicspac-
without any Judgingfrom Kdley's er regions were generated
specific reasons. forL. haenkei fOllowing
diagnosticcharacters, L. insignisseems quitedis- the proceduresof Kress et aL (2002), including
tinct. Nevertheless,no reconsideration has been DNA extraction, amplification, and sequencing.
made to date. The third species L. auriculata Phylogeneticanalyses fo11owedthesame pro-
appeared in Elmer (1939).
But the article is just
a cedures as Kress et al. (2002).
Maximum parsimo-
kindof a personallist
of unpublished naJnes. Elmer ny ana]yses of the ITS and matK data
sequence

(1939)
stated,
"The
fol]owing
numbers of my col- were conducted using PAUP"4.0 (Swoffbrd 1998)
lectionwith unpub]ished new names be con-
should with unweighted characters and 500 random-
sidered publishedspecific names as indicated
below sequence-addition replicates, saving all shortest

with theirauthors," "17977-Leptosolena auricula- trees under TBR Branch Swapping, STEEPEST
ta isLeptosolenahaenkeiPresl."
Therefore
we will DESCENT off, MULTREES on, COLLAPSE

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AprJl2005 FUNAKOSHI et aiRedigcoveryofLeptosolenahaenkeiZingiberaceae 43

wwkl iilee\ aj･lli

lltw'ei,
}esee},lt-tamu/#

mamaE
'lwai pm--e
imum.
,lmaelllmalww,illvat
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ne

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aspmer;
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,'{,ilstkl
:-}ttMA.':ilX,l,,,,Iikgevavampnl// .

, l'wwllllIlwwlll,,l

leei}XleeII
,, mekl

FJG 1 The leLtotypcot LeptosolenahaenkeiC Presl(T HaenkE s n bheet no 335771 1832A, pR) wtth K BPrcsl s handwnting
Cupperportionidentified byB Skeedopoloya pR)

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44 APG Vbl.56

branches if maximum length is zero. Bootstrap

analyses were conducted RAUP'4.0 with ten
using First
of al1,
Ridley'sdescription
incalyx length
random addition replicates,TBR branchswapping, of Leptosolena haenkeias 1.5cm is very much
for1OO bootstrap replicates. The datasets t'or
each likelyto be a mistake of 1.5inches,becausein
gene region were analyzed separately and then,fo1- Presl'soriginal descriptien itwas describedas
lowing the total evidence approach for multiple "sesquipollicaris"
which means 1,5 inches.
datasets, combined. Haenke's specimens liebetween Ridley'sdescription
of L, haenkeiand L, insignis.
Resultsand Discussion Inthe typesof Leptosolenainsignis(holotype,
K, non vidi; isotype,Ny, digital
image seen; p, dig-
7Zixonomic comparison between Leptosolena italimage seen; slNG!; us, digitalimage seen; type
haenkei and L. insignis location describedin Turner (2000)),
distinct
We first examined the concordance of character pedicelscould be recognized buttoo short to mea-
figuresinthe text of Presl's(1827) original descrip- sure theirlengthin the isotypeat us, but they were

tionofLqptosolena haenkei,attached illustrationin obvious in the isotypesat p, slNG, and Ny, [E]he
Presl(1827)
with notes as drawn in actuai specimen measured characters are: calyx 7,2-7.8 cm long
size, and our measurement (five
in Haenke's
flowersmeasured, siNG), corolla tube 12.7 cm
original

materials (four
sheets) depositedat PR and PRC. (1) (twoflowers measured, slNG), leaves 31 × 2 cm
In pedicel,written as pediceledin Presl'stext, (NY), 30 1.6¢ m (s]NG), 25 2 cm (us). All leaves × ×

pediceledinillustration,
pediceledinoriginal mate- were in condition foldedin halffrom
mounted

rials we measured. (2)In calyx

length,3.8cm in midrib, buttheleafwidth isalmost doublethe figure
text,4,5cm 4.2- 5 cm in original
in illustration, which Ridleydescribed. Then thediagnosticchar-

material (1Oflowersmeasured), (3)In corolla tube acters from our recent collection from Northern
8 - 9 cm
length,7.5cm intext,9,1 cm inillustration, Luzon, Philippineswere measured (five
flowers
inoriginal material (five
flowersmeasured). (4)In from three plants preserved in spirits were mea-
corolla lobelength, 1,5cm intext,1.4cm inillus- sured, leaveswere separately collected from the
tration.(5)In anther crest, divergedin text,not middle of the pseudostems); calyx 7.3-7,8cm,
developedin illustration. (6)In leaves,25 - 27.5 corolla tube 11,O-11,5 cm, leaves 42-45× 3-3,2cm.
cm × 2,5cm intext,20 - 31 cm × 2 - 2.8cm inorig- Wb confirrned thepresence of distinctpedicel(4mm
inalmaterial. long)and a little developedanther crest.
Accordingto Ridley(1909), the differences Inthe other specimens examined, calyx length
betweenthe two species are as follows. are 6.5cm (L2znoverhergh s.n., BR); 6.0crn(Santos
5674a, L); 6.8cm (Steiner 2198, L); 5.5cm (Fox
230, GH). Corolla tube are (Silanoverbergh
10,7cm
Flowers pediceled, calyx 1.5 cm long,corolla tube s.n., BR); 10.6cm (Santos
5674a,L); 10,9cm(Fox
7 cm long,corolla lobe 1,2cm long,anther crest not 230, GH).
""''"'
prolonged, leaveslanceolate (30× 2,2cm) Our recent collection matches well with neither
-･･--････--･･--･･-----･･--･t･･････--Leptosotenahaenkei
of these two species'descriptions, or rather with a

Flowerssessile, calyx 6 cm long,corolla tube 12 cm mixture ef these two descriptions; which means

Iong,corolla lobe 2 cm long,anther crest 1arge, thatLq)tosolena haenkei, L. insignis,

and our recent

leaveslinear (25× 1 cm) "''""''""''"L.

insignis collection are conspecific, resulting inL. insignisas
latersynonym. The character figures derivedfirom

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herbariumspecimens of Leptosolena 2 (6): 23 (1889);

other than
RidL, Philipp.J. Sci.C. Bot.
types which liein-between support thatL. 4: 181 (1909);haenkei
Merr, Enum. Philipp.Fl,Pl,1:
and L insignisare conspecific. Tb avoid confu- 236 (1925); Larsen et al., in Kubitzki,Fam.
sion, we providea more accurate descriptionof L. Gen. Vasc.Pl.4: 492 (1998). tep- =Aipinia

haenkeibasedon our recent freshcollection togeth- tosotenia K. Schum.,Pfianzeur.IV 46: 312

er with character infbrmationobserved inall exam- (1904), with incorrect author citation as (C.

ined herl]ariumspecimens. Presl) K. Schum. non A. haenkeiC, Presl

(1832). -Lectotype: Philippines, Luzon, Z
711ie lectotyl)ijicationofLeptosolena haenkeiC. Haenke s.n. (pR,sheet no.335771-1832A
PreslAs [Fig.1], iso-:pR, sheet nos. 335771-1832B,
fbr Haenke's specimens were distributed to 335771-1832C;pRc). (here designated)
about 20 hefoaria(Stearn 1973)which includesB, Leptosolena insignisRidl., Publ.Bur.Sci.Govt.
BM, BR, F, G, GH, GOET, K, L, LE, M, Ny, and W Lal).35: 84 (1906); Ridl.,Philpp.J.Sci.C.
(Sko6dopolova 1995). For inquiry fbr locating Bot. 181 (1909);
4: Elmer., Leafl. Philipp. Bot.
Haenke's original materials, only negative respons- 8:2907 (1915). Luzon, Benguet Prov.,
-7)Tpe:
es were derived from B, BR, F, GH, L, M, Ny, and w. [[iwinPeaks,A. D. E. ELmer 6428 (K, non vidi;

As ferHaenke'sspecimens of Leptosotenuhaenkei, Ny [digital image!], P [digital irnage!],sLNG!, us

Schumann (1904) stated that specimen has"type
[digitalimage!]).
probablybeen lostbecause it doesn'texist any- Leptosotena auriculata Etmer,Leafl. Phi1,Bot. 10:
more in German Universityor Bohemian Museum 3808.(1939)., nom. nud. -Specimen cited: cul-
at Prague." But Haenke'scollection of L. haenkei tivated at Los Banos, Laguna Prov. Luzon A.

are currently extant at PRC (Charles

University D. E EImer17997(F, non yidi; L, non vidi; Ny,
which was formerlycalled German University; one non vidi; w [digital
image!]).
sheet) and PR (threesheets) even though Schumann

actually confirmed some other Zingiberaceaespec- Perennial,evergreen herb.Pseudostemserect,

imens (e.g. Aipinia moltis C. Presl,Aipiniabne- not droopingtoward the top, 280-330 cm long.
vilahris C. Presl,IZblowratia elegans C. Presl)at Rhizomeselongate, with ca. 20 cm intervalbetween
PRC judging from his determinavitslips as neighboring pseudostems. Plane ofdtstichy of leaves
"Bearbeitet
furdasPfianzenreich" (Tlr;eated
forthe isperpendicularto rhizome. Leavesglal)rous, sessile,

Pflanzenreich).Nevenheless, the misunderstand- linear, numerous, ca. 66-75 leafbladesper pseu-

ing of the typeas lostas suspected inBurtt& Smith dostem at floweringseason, base cuneate-round-
(1972)who believed Schumann's note prevailed ed, apex caudate-attenuate, 42-45 × 3-3.2cm in the
unti1 recently. AlthoughPresldidn't
cite any par- middleof pseudostem. Ligutesglabrous, as an erect

ticular specimens inhisprotologueof L. haenkeiin pairof auricles, oval, rounded at tips, 7 mm long.
Reliquiae Haenkeanae, Haenke's extant collection of 1njiorescence glal)rous,paniculate, 7-11cm long,ter-
thespecies at PR and PRC should be considered as minal on thepseudostem, with 2-4brown persistent

elements of original material. Here we select the inflorescence bracts.Lower cincinni have 5-6flow-
most well-represented specimen as the Iectotype ers with distinct pedicelswhich are 4 mm long.
of L.haenkei(Fig. 1). Floral bracts subtending cincinni, and bracteoles

absent. CZilyxtubular, irregularly

2-3 lobed shortly,
Leptosolenahaenkei C. Presl,Reliq. Haenk. 1: split deeperon one side, 4.2-7.8cm long.Cbrolla
111,t.18 (1827);
Petersen,in Engl.,Pflanzenf. tubes long and narrow, white, 8.0-12.7
cm long.

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46 APG Xlol.56

RG. 2.A: Infiorescence haenkeii fuIIbloom (at

of [eptosolen[i 2000 hT).B: Hal)itat
ofL haenkoi.C: Upper ponion ef fiowcdngpseu-
dostemofL haenkei i itshabitat.
D: Whole plantofL lzctenkei.
E: InfructescenceofL haenkei.A, C, D (Fbenakoshi& Cb 2CV6),
E(Fun`rkoshi2025).

itwith Alpiniaeiegans, A. Iuteocarpa,

and thetwo 95%).
species of Vcmovetherghia.However, strong support The themolecular analyses suggest
results ef

(bootstrap
value 93%) isprovidedforuniting itin
= thatLeptosolena evolved ina clade with other taxa
a clade with Etlingeraand Hbrnstedtia
as well as insubfamily Alpinioideae mibeAlpinieaenow dis-
with several species ofAmomum (bootstrap
value = tributedin thePhilippines (e.g.,
V2inoverberghia

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of Le7)tosolena
haenkei,Zingiberaceae 47

t/''/1t'ttt't
'''tt't{ttt
'./.,'tttt''
'J.rtt'/tt.-.tttt'･･I-
'iti.4.,.,gt/ttt'':'t""tl'f'':ri'''r-

･r

ll,''
1i'l

' ''''
'' ''-v''' E
'tt,, ts

A.1' t'/"'',..#''
'"l1' ! .:
't/t4tttt 'it'/ttt'
3cm

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.,t=Ab..-:.,,,/t:.ttt
''

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.. ..SA-
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ti,g
lo, '.,.
`z J
H
FIG,3.heptosolenahaenkeiC. Presl. A-E H-I (Funakoshi & Cb 2006),G, J (Funakoshi 2025).A: flower, dissected,B: calyx. C: close-
up of labellumbase.D: fiower,side view. E: flower,
frentalview. F: anther and anther crest G: capsu]e. H: nectary and ovary. I:
infiorescence.
J:transverse section of capsule.

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CorollaZobesoblong-ovate, apex obtuse, 2 cm Otherspecimensexaminedi PHILIPPINES, Supan,

long.Lateralstaminode petaloids, obcordate, apex Meuntain Prov,,Luzon, Jun,4, 1911,ML ltitnoverbergh
s.n. (BR);Bauan - Mt. [Ihbuan, Cagayan Prov.,Luzon,
of thetwo lobessubacute, 7 mm long.Labeitum
May, 1929, M. Ramos 77052 (Ny,slNG); Villar, ML
oval, erese-crisped, fragile,recurved, white with
Pinatubo,Zambales Prov.,Luzon, alt. 350 m, May 18,
no nectary guideobserved to human naked eyes, 194g,R. B, Fox 230 (GH, pNH); SitioDandanak, Banio
4.7-5.1 × 3.5-4.0 cm and palatepubescent,Filament Bana-aw,Municip,Besao,Bontoc Subprev,, Meuntain
almostunable to be recognized. Anther8 mm long Prov.,Luzon, alt, 2600 ft,May 5, 1953,J. U Sant'os
with a little developed anther crest. Stigma cup- 5674a (L,puH); road Bontoc -Mt Data,MountainProv.,

shaped, with hairsalong the edge of the hollow.

May 14, 1961, M. L. Steiner2198 (L);Brgy Alab,
Municip. Bontoc, Mountain Prov.,Luzon, alt. 1100 m,
OvaT:),3-locular,axial p]acentation, 8 × 5 mm,
May 3,2oo2,HL Funakoshi& L. L Cb 2006 (AAu,
E, puH,
Cbpsulesglabrous, oblong, slightlytrigonous,con- Kyo, us); Brgy Balili,Municip. Bontoc, Mountain Prov.,
taining numerous persistent alt. 940
arillate seeds, with m, Sept.29,2002,H. Funakoshi2025 (E,
GH,
calyx at apex, 4.6-5.1× 2.0-2,5cm excluding per- PUH, KYO, L)

sistent calyx length,

Seedssubglobose, irregularin
shape, ca. 2 mm in diameter. PhylogeneticplacementofLeptosolena
haenkei in
PHILTPPINES, NorthernLuzon
Distribution: theZingiberaceae
(Benguet,
Cagayan,IlocosNorte,La Union,Moun- I7S - The analysis of the ITS sequence dataresult-
tain Prov., Nueva Vizcaya, Zambales) (additioned in six equally parsimonious trees of 743 steps

made [1925]);
to Merrill alt, 300-1300m. (consistency
index [CI] =
O.479;retention index
Uabitat and Ecology: Abundant on sunny [RI] = O,750; rescaled consistency index [RC] =

rocky s16pes where even dominantPinus kesiya O.359;Fig. 4).Leptosolena isstrongly supported

forestare unable to be sustained (Fig.

2B). This (bootstrap
value 96%) as a member = of a clade

species ischaracterized by thenocturnal anthesis; its containing species ofAipinia, l,Zinoverberghia,

label]umstarts expanding a few hoursbefbresunset, Etlingera, and Hbrnstedtiaalthough there isonly
This phenomenon frequently
' led the incorrect poor bootstrapsupport foritsclosest relatives with-

descriptionof the fiower. In Ridley (1906),itwas inthis clade,

described as "Lip
fleshy,elongate, 1 i'nch long," matK - The analysis of the matK region (cod-
but actually thelabellumlooks likea wrinkled frag- ing and noncoding) resulted in 120 equally parsi-
ilepaper reaching 5 cm longafterca,itsfu11expan- monious (CI O.703;RI O,882;
treesof 518 steps = =

sion after sunset (Fig.2A), The floweralso emits a RC O,6230;Fig.5).Leptosolenaisstrongly sup-

=

sweet fi;agrance duringitsanthesis. Those charac- ported (bootstrap value 93%) as sister to Alpinta
=

teristicstogetherwith snow white flowercolor, no vittata but only has weak bootstrap support as a
visible nectary guide,long(ca. 11 cm) and narrow member of the clade made-up ofAipinia elegans, A.
corolla tube and almost no filament unlike tvtteocarpa,and the two species of ilanoverberghia.
Heclychium might imply thepollination
by special- Only poor support isprovided foruniting itin a
istlikenon-hovering moths with a very longpro- c]ade with Etlingera and Hbrnstedtia.

boscis.Itisalso worth to mention about the unique Combined data set - The analysis of the com-
fruit;
relatively big ftuits (Fig.2E, ca, 5 × 2,5cm) binedITS and matK sequence data resulted in 64
contain numerous arillate seeds which taste sweet parsimonioustrees of 1,286steps (CI
equatly =

and This probablymeans

sour. seed dispersal
by O,564;RI O.803;RC O.453;Fig.6).Leptosolena
= =:

fruitbats,but the conclusion must await further isweakly supported (bootstrap yalue 59%) as =

field
observation. sister to Aipinia
vittata with poor support inuniting

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Ahamomum angustifoiium
Aframomum sceptrum

Aframornum denieltii
Aframorrium species #1
Aframomurn species #2
Reneatmia alpinia
Reneelmia thyrsoidea
Reneatmia cernua
Renealmia species #1
Renealmia species #2
Renealmia battenbergiana
Amomum glabrum
Amomum longipetiolatum
Elettariopsis
kenbyi
Etettariopsis
stenosiphon
Etettariopsis
species
Paramomumpetaioideum
Alpinia carolinensis
Alpiniaelegans
8311 Alpiniaiuteocarpa
Vbnoverberghia sepulchrei

Vanovetheighiasasakiana
Alpiniavittata
Leptoselena haenkei
Etlingera etatior
Etlingere yunnanensis
Homstedtia hainanensis
Etlingera iittoralis
Amomum vitiosum

Amomurn species

A/pinia biepharocalyx
AipiniainteiTnedia
Aipiniapumila
Plagiostachysspecies #1
Plagiostachys species #2
Aipinia catcarata
100 Aipinia oM'cinarum
13
Aipiniafoxworthyi
Siliquamomum tonkinense
But:bidgeanitida
Burhidgea schizocheiia
Pleuranthodiumfioribundum
Pleuranthodiumsch/echteri
Riedetia species
Siemanthus siJiquosus
Aipinia conchigera
Alpinia galanga
famijia fiagetJaris
Siphonochiiusaethiopicus
Siphonochiius decorus
Siphonochiiuskirkii

FjG. 4.0ne of the six

equally parsimonioustrees of subfamily Alpinieideaeinthe analysis of the ITS sequence dataClength743; =

consistency index O.479 excluding uninformative

= churactcrs; rctcntion index O.750;and rescaled consistency index
= O.3S9) =

showing branch Lengths(above the line)and bootstrapvalues (below the lineif>- 50%). Asterisks
indicatenodes that collapse
in the strict consensus tree.

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24 Ahamomum angustifolium

Ahamomum daniellii
Attamomum sceptrum

Afiianiomum species #1
Aframomum species #2
Renea/mia aipinia
Renea/mia thyrsoidea
Reneairr?is species #1
Reneaimia cernua
Renea/mia battenbetyiana
Reneatmia species #2
Amomum gtabrum
Amomum iongipetiotatum
evettariopsis kembMi
Elettariopsis stenosiphon
Elettariopsis species
Paramomum petaloideum
Alpinia conchigera
Aipiniagaianga
Aipinia bJepharocal)or
942 Aipiniacatcarata
Aipinia intermedia
Alpinie purnita
AipiniaoMcinarum
Atpiniafoxwotthyi
Piagiostachys species #l
Piagiostachys species #2
Atpinia eiegans
Atpinia luteocarpa
Vlanovet:betyhia sepuichrei
Venoverberghia sasakiena
Alpinia vittata
Leptosofena haenkei
Etiingeraelatior
Etiingerayunnanensis
Hornstedtiahainanensis
Etiingeraiittoralis
Amomum viltosurn
Amomum species
Aipiniacarolinensis
Burbidgea nitida
Burbidgea schizocheila

Pieuranthodiumfiorihundum
Pieuranthodiumschlechteri
Riedeliaspecies
Siemanthussiiiquosus
Siliquamomum tonkinense
tamijia fiageileris
Siphonochilusaethiopicus
Siphonochitus decorus
Siphonochiluskirkii

FiG,5. 0ne the 120 equally parsimonious

ef trees of subfamily Alpinioideae in the analysisof the mati( region (codingand noncoding)
sequence data(length518;consistency index O.703 exc]uding uninformative
= = characters; retention index O,882;
=
and rescaled
consistency index= O.620)showing branch]engths(above the line)
and bootstrap values (below theHne if) 50%), Asterisks indi-

catc nodes that collapse in the strict consensus tree.

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7Ahamomumangustifotium
Aftamomumsceptrum

Afrumomum danieltii
Afrumomum species #1
Ahamomum species #2
Reneafmia aipinia

Reneatmia thyrsoidea
Renealmia species #i
Renealmia cernua
Renealrnia battenbergiana
Reneatmia spedes #2
Amomum gtabrum
Amomum Jongipetiotatum
evettariopsis
kerbyi
blettariopsis stenosiphon
evettariopsis
species
Raramomum petaloideum
AVpinia conchigere
Alpiniagaianga
Alpinia blepharocalyx
Alpinia intennedia
Alpiniapumila
Alpinia calcarata

AlpiniaoM'cinarum
Aipiniafoxworthyi
Ptagiostachysspecies #1
Plagiostachys species #2
Aipinia carolinensis
Aipiniaeiegans
9212 Aipinia luteocarpa
Vlanovetherghiasepulchrei
VZenoveibetghia sasakiana
Aipiniavittata
Leptosolena haenkei
Etiingera eletior

Etiingerayunnanensis
'Hornstedtia
hainanensis
Etlingera Jittoratis
Amomum villosurr]
Amomum species

Siliquamomum tonkinense
Burbictgea nitida
Bunbidgea schizocheila
Pleuranthodium fioribundum
Pleuranthodiumschfechteri
Riedelia species

Siamanthus siliquosus
lamijiafiagellaris
Siphonochilusaethiopicus
Siphonochilus decorus
Siphonochilus kirkii

FIG,6. 0ne ofthe 64 equa]ly parsimonioustrees of subfami[y Alpinioidcae in thc analysis of the combined ITS and matK region
sequcncc data(Iength1,286;consistency index O.564excluding uninformative characteis; retention index O.803;and rescaled
= = =

consistency index O.453)showing branch lengths(abovethe line)and bootstrap

=
vul ues (belowthclineif>- 50%). Asterisksindi-
cate nodes that collapse in the strict consensus tree.

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TABLE1. Listof species used inthe phylogenetic theZingiberaceae.

analyses of Nbucher number, location,
eountry of origin, and
GenBank accessjon number of gene sequencesprovided.
are

[[bxon name Nbucherinformation: Ceuntry of origin GenBank GenBank

co]lector, nurnber, and accesslon accession

herbariumlocation number for ]TS number formatK

sequences sequences

Leptosolena haenkei C.PreslFunakoshi & Co 2006, US Philippines AY742331 irv742390

seputchrei, Aipinialuteocaipa,and A, elegans) and extremelylongand narrow corolla tube,the lackof

Oceania(A.vittata) and isalso related to species of floralbractsand bracteoles, and especially the
the genera Amomt{m, Etlingera, and Hbrnstedtia Iateral
petaloid staminodes, which are exceptional in
(Kress et al. 2002).Contrary to the notion ofBen- subfamily Alpinioideae,make Leptosotena a read-

tham (1883) and Burtt& Smith(1972), Leptosolena ilyrecognizable genusinZingiberaceae.

is only distantly related to the Bornean genus The phylogenetic placement of Lqptosolena
Burbic(gea in tribeRiedelieae, No evidence sup- in a clade containing species ef the highlypoly-
portsRidley's(1909) contention that this genus is phyleticgenus Aipiniaisproblematic fordelimiting
closely related to Heclychium placed in subfamily genericboundaryinthisnibe.Although Leptosolena
ZingiberoideaeHaask, (Kress
et aZ, 2002), is morphologically distinct,
itis yetundetermined
In our ana]ysis the genusLqptosotenaissister how species can be combined into monophyletic
to Aipiniavittata, but isquiteunlike itsclosest rel- genera in these clades. Further analyses of addi-

ative in morphological characters; viz. 11 cm vs, 2 tionaltaxa using rnolecuJar characters and more

em incorolla tube length,no floralbractand bracte- in-depth studies of morphological character variation

ole vs. conspicuous bractand tubular bracteole,a are warranted beforenew generic boundariescan be
well-developed labellum
vs. an inconspicuouslabel- established.
lum that isalmost thesame size as thecorolla lobes.
Alpinia elegans andA, The authors are gratefulto LeonardoCo (puH)
luteocarpa,both belonging fbrgiving
to sect. Kotowratiaendemic to the Philippines fullfacilities forthe fieldsurvey in the Philippines. We
greatly appreciate a deep insighton the technical term
(Smith (1990), Funakoshi unpublished data),have
given from Kai Larsen (AAu)and Mark Newman (E),
some characteristics incommon with Leptosotenain
We wish to express our gratitude to ShutaroNishihara,
their nocturnal anthesis and relatively largefruits Mutsuko Nakajima, and Ida Lopez for
preparingFig. 2,
(Funakoshi Fig.3, and Figs.4-6,respectively. Our
"sweet
with and souT" arillate seeds sincere thanks also

unpublished data).iinnoverberghia, another inter- go to the directorsand curators of B, BO, BR, F, GH, L, M,

esting representative of the Zingiberaceae in the NY, P, PNH, PR, PRC, puH, SINa, us, and w fbrthe permission
to examine the specimens or forthe search forlocating
Philippines, isquitedifferent from Leptosolenain
specimens or for providing Web sites to see the digital
the conspicuous bractssubtendjng one flower,the images of type specimens. We acknowledge JinMurata
deeplybilobedlabellumbasallyconnate to the (TI), Hideki Nakai, HidetoshiNagamasu (Kyo),and
corolla lobes,and the channeled filamentenclosing Masanao Honda fbrprovidinginvaluable comments on

the style. Otherspecies (Etlingera elatioi; E,yun- the manuscript.

nanensis, Hor:nstedtia hainanensis) inc]udedinthe

same clade with Leptosotena differ inpossessing References
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Hooker. Genera Plantarum.Vbl.3.p.647.Lovell
and i/kenoverbeighia.As mentioned earlier, the
Reeve,London,

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April 2005 FUNAKOSHI et al.r Rediscoveryof Leptesotenahaenkei,Zingiberaceae 53

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Prantl(eds.) Die'Naturlichen Pflanzenfumilien, 1st somal RNA genes for phylogenies. in: Innis,M. A,,
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ReceivedOctober15,2004raccepted lanuary13,2005

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