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ISSN 1346-7565
Acta Phytotax,GeobeL 58 (1):
19-32 (2007)
Myxochinmys(Zingiberaceae),
a new
genus
from Borneo
ATSUKO
TAKANoiand
HIDETOSHI
NAGAMASU2
iMtiseum
Ri,ogo 669-154a Jinpan;
Activities,
ilyogo,6-chome,}Zryoigaoka,
Skencla,
andHuman
ofNdture
2111te
UhiversityMitseum, K),oto
UhiversiijlSZik}'o,
606-8501,Jbpan
K)?oto
K),oto
A new genusand species, filyxochtanzys
muilerensis
from CentralKalimantan,
Borneo
(Zingiberaceae)
infiorescences
with
isdescribed,
Thisnew species ischaracterized by indistinct
congested
pseudostems;
from
spirally arranged bracts;
copious, highlyviscous, axjllary-pooled
mucus
secreted
glandson
imbricatebracts;and fiowerswith a concave labellum,versatile anthers, and a grooveforrned
by the inner
surface of thecorolla tube to envelope and holdthe style. Parallel
orientation ofthe planeofdistichy of
the leafyshoots, well-developcd
staminodes,
and labellumnot cennate
with
the filament
suggest
that
Adyxochlanrysbelongsto the tribe Zingibereae,Among the generain Zingibereae,Adyxoehlanrys issimilarto Staphochlanrys
inthat itlacksa distinct
itisdistinct
from
pseudostem and spiral inflorescence;
Sbaphochlaiays,
however,inthatithas a groovewithin the corolla tube, concave labellum
and versatile
anthers, Phylogenetic
analyses
basedon DNA sequences
of the nuclear
internal
transcribedspacer (ITS)
K
and
maturase
and
Sbaphoehlamps.
The
ofthe'chloreplast
(matlO
similarities and
DNA
show
similarity
with
Distichochlanv]s,Adyxochlanrys,
differences
betweenA()xeehldop;s
and
related
generaare dis-
cussed.
Key words: Borneo,toxochlamys,
mullerensis,
toxochlanrys
Sbaphoehlantys,Zingiberaceae,Zingibereae
The Zingiberaceaecomprise
ilyof53
and
over
genera
1200
a
fampantropical
species
et al.
(Kress
molecular
phylegcny,ncw genus,
Alpinieae),and ZingiberoideaeHaask.
the remaining
Currentstudies
genera),
(containing
on
the mol-
diversity
is southern and
ecular phylogenies
of the Alpinioideae and some
southeastern Asia(Larsen
et al. 1998).
TraditionallMlarge genera such as Aipinia,Amomum,
and
fburtribes (Alpinieae
Meisn.,Globbeae Meisn., Etlingera
2004,Xia et aL 2004,Kresset
(Pedersen
Hedychieae Petersen, and Zingibereae Burtt &
al, 2005)suggest thatthe Zingiberaceae
are monoOlatubji)
havebeenrecognized inthe familybased phyletic,
butrequire redefined genericboundaries,
et al.
especially
fbrAipinia and Amomum, which are
on fioraland vegetative characters (Dahlgren
2002) with
a center of
1985;Larsen et
al.
199g).Kress et al.
recent(2002)
In Zingiberoideae,
Williams
polyphyletic,
showed
that Globbeae are mono(2004)
apparently
lyproposed a new classification ofthe Zingiberaceaeet al.
basedon a molecular phylogeneticanalysis, They
phyletic and sister to Zingibereae; however, the
recognized
fbursubfamilies: Siphonochiloideae
W, generic1irnjts
intheZingibereae
remain unresolved.
J. Kress (containing
only Siphonochilus),[lamijiIn 2004, we explored Muller Range, Central
oideae W, J. Kress (containing
only 7Zimijia), Kalimantan, Indonesia and found an interesting
of
Alpinioideae
Link (containing
most of the fbrmer
orientation ofthe planeof distichy
ginger.Parallel
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APG
the leafy shoots, well-developed
staminodes,
and
labellum
not connate to the filament
suggest thatthe
plantbelongsto the tribeZingibereae,but comparative morphological
studies
and molecular
provideevidence
against
phylogenetic
the placement
of
Xlo1.
58
54eC. Amplified productswere
Microspin S-300 HR columns
purifiedusing
(Amersham
Biosciences, Buckinghamshire,England,UK).
Sequencingwas perfbrmedusjng ABI Prism(tm)
Big Dye Terminator Cycle Sequencing Ready
Reaction Kit (Applied
Biosystems,FosterCity,
Califbrnia,
USA), Sequencingprimersincludedthe
thisplantwithin any ofgenera inthis tribe.Thus, a
new
and new species, M
genus, ILd),xochlantys,
mullerensis,
isdescribed
here. In accordance with
amplification
primers as well as ITS2K (Rangsiniji
Article42 of the InternationalCode of Botanical et al. 2000)and ITS3P (M611er
& Cronk 1997) fbr
Nomenclature(McNeill
et aL 2006),
the names of
the ITS region and m5R, mSP2R,
m5fa, m8R,
the new genus and species are simultaneously
datedusing the description
provided below.
vali-
Materialsand Methods
matK8R
from Kress
(all
2002), matK8
et aL
(Steele
& Vilgalys1994),and [EA05R (Takano
& Okada
2002) fbrthe matK region. The productswere
cleaned
using
the manufacturer's
and were
protocol,
run on an ABI Prism'M 310 GeneticAnalyzer
Biosystems Japan,Tokyo, Japan),Raw
(Applied
Descriptionofthe
species
The species description
isbasedon studies of
plants in the fieldand in the herbaria of BO, E,
HYO, K, and KYO, The plant material studied
includes
pressed and driedspecimens, spirit-fixed
floral
material, and silica-gel dried
tissueformolecular phylogenetic
analysis.
sequences
were
assembled
and
edited
using
the
BioEditver. 5.0.9 software (Hall1999). DNA
sequences were aligned by multiple
alignments
using the CLUSTAIIWI ,83 computer
software package with
defaultsettings
et al.
(Thompson
1994).
Pdylageneticanalyses
Pbylogenetic
analysis
For the DNA
somal
DNA
of
using
DAC4
sequence
clata
analysis, we used nuclear ribothe internaltranscribed
spacer
ITS1, 5,8S,and
(ITSs;
of the chloroplast
(matl()
regions
The ITS dataset
was analyzed
bothseparately
and in combination
with the matK
dataset.
To test
dataincongruence,
the incongruence lengthdifTerence (ILD)
test(Fanis
et aL 1995)was implemented as the partition
homogeneity test option in
PAUP" with 100 replicates foreach of two parti-
ITS2)and maturase
K
DNA. The ITS and
matK
sequences
inA(lyxochlanrys
were generatedby
the fo11owing
method
and deposited
in theDNA
tions.
Data Bank of Japan (DDBJ).
The sequences
of
Maximum parsimony (MP)was amalyzed using
other taxawere obtained fromGenBank (AppendixPAUP" ver. 4,O.
beta10 (Swoffbrd
2003). Heuristic
rlbtal
1).
with
SIMPLE addition
genomicDNAs were extracted fromsilica- searches were conducted
driedtissuesby using the modified C[IIABmethod
tree bisection-reconnection
(TBR)branchswap&
Doyle
1987,
Takano
&
Okada
2002).
(Doyle
ping,and the MULPARS options, Only basesubITS sequences were amplified using the ITS5P and
stitutionswere treatedas characters fbrphylogeny
ITS8P prirners(M611er
& Cronk 1997).The chlororeconstmction,
Allthe gaps were treated as missing
,
plast mati(;
region
was
amplified
using trnKIF
(Manos& Steele1997)
and trnK2621 (Liston
&
Kadereit 1995).All amplifications were perfbrmed
using
TaKaRa Ex [ktqTM
Bio Inc.,Otsu,
([lakara
Shiga,Japan),at annealing
temperatures
of
50"to
values.
Support forbranches was
bootstrap analysis
estimated
using
1000 replications
1985) by a heuristic
search using SIM(Felsenstein
PLE addition, NNT branch swapping,
and the
MULIIARS options, Three species ofSiphonochitus
with
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were
TAKANO
selected
et al. 2002).
(Kress
the outgroup
as
& NAGAMASU:
21
tauroehtam.v,y,
gcn.nov,
duobus;ovarium
A Bayesian
analysis
was
eonducted
v,
Central Kalimantan. Murung
using
Raya
MrBayes
basali.
placentatione
uniloculare,
7]tpus,INDONESIA,
District, behind Tumbang
Naan
(O'09'03"S,
3,1.2(Huelsenbeck
& Rohnquist 2001,
from Sungai(River)
Lapangan,a
Rohnquist & Huelsenbeck 2003). The bestfitting
branch of Sungai Joloi(OO11'OO"N
113031'59"E),ca,
fbrBayesian analysis was selectsubstitution model
200 m from Muara Sopan (ee12'22"N
113e31'23"E),
ed using a series of likelihood
ratio testsas imple- December 21, 2004, H. Okada, H. Ndgamasu, H.
mented
in Modeltest v. 3.7 (Posada
& Crandall 7lsukLu/a,
A. lahano,&A. AJdiki,
K7L671 (HoloBO; isoHYO, KYO),
1998).We ran over one million generations
of fbur
113045'04"E)
simultaneous
cold,
(one
en
route
three heated)
Matkov chains
Rhizomatous herbs.Rhizome slender, elonby using MrBayes y, 3.1.2until the average standard
gate,Leafy shoots close together. Leaves sessile, 3deviation
of split firequencies
became less 9,distichous,
(ASDSF)
to rhizome.
Planeof leavesparallel
than O.O05.The tree from every 100th generation
Leafsheathopen to thcbase,17.5cm long,ca. 1 cm
was retained (excluding
a quarter
of 1OO/total
gen- wide; ligule,
5 mni long,Leafblades narrowly oboerations), and a 50% majority
rule consensus
tree vatc, to 50 cm long,ca. 7 cm wide, basecuneate,
was constructed
from these. For each ITS and the apex acute, upper surfhce sparsely hairy,uniformly
combined
dataset,
the logstabilized before50 millower surface densety hairy,unifbrmly
glandular;
liongenerations
in the Bayesian analysis (1,OOO,OOO
Inflorescence
terminal on a leafy
shoot, 4glandular.
forthe ITS dataset
and 4,500,OOO fbrthe combined
15 cm long,fiowering
from the baseto apex; scape
dataset),
The results of theBayesiananalyses are
2.5-4 cm long.Bracts spirally arranged, imbricate,
reported
as the posterior probabilities (PP;
ca. 5 cm long,
ca. 1.3cm wide,
concave, uniforrnly
Huelsenbeck & Rohnquist 2001),which are equal to
puberuleus, with many
glands,blood red, fi11ed
the percentage
is resolved.
trees sampled
of
Only PP
when
scores
a
excess
given clade
of 509,6are
with
to bracts,
ca. 3.5cm long,ca. 1.3
keeled,opposite
inour
shown
trees,
cm
The
result
of exploratory analyses
diMcult-to-align
regions
thatexcluded
wide,
resolved
(not
117
(ITS,
Calyx tubular, ca, 2.5 cm
ment
dataset excluding
within the ITSs,
regions
2
with
results
from the
ambiguous
align-
Results
ca.
mullerensis
gen.et
1,2)
(Figs.
1.5cm
wide,
concave,
gin. Lateralstaminodes
sp. nov.
ca.
Herba
fbliadisticha
parallelead rhizoma.
fbliatoterminalis, e basi ad
dispositae,
fioremsinfiorens;
bracteae
spiraliter
rhizomatesa;
Infiorescentiain surculo
apicem
mm
3 mm
wide,
subtendens,
mucum
gelatinum copiosum
white.
long,with
and e:Tveloping
long. Ovary
gularem
3 unequal
teeth, glabrous,
Corollatube ca. 4.7 cm long,thick, with a groove
fbrmed by innersurface enveloping style; lateral
petalsovate, white with red apex, ca. 2 cm long,ca.
O.8cm wide,slightly hooded;dorsal
petalovate, ca.
2,1 cm long, ca, O.9 cm wide, slightly hooded,
beaked,totally glal)rous.
Labe]lum ca. 2.8cm long,
A. [[lakano& Nagam.
Myxochlamys
or
topologies;there-
pairs)produced
fbre,in thispaper,we present
only
ofthe
base.Flowerssolitary.
long,split along one side
to
open
glabrous,
shown)
to base, apex
more
analyses
Bracteoles2-
transparent mucilage,
colorless,
a
pinkishmarobovate,
ca. 5 mm
long,
Antherwhite; filament
ca. 5
groove
style;
ca,
white
5
mm
connected
with
to corolla tube
thecae versatile; spur ca.
6 mm
long, glabrous,
unilocular,
secer-
basal,rudimentary
septa
as
placentation
present
bracteolaebracteisoppositae,
bicarinatae,
ad
longitudinal
ridges
inside
ovary. Epigynousglands
basinapertae. Florescorollae tubo et filamento
intraproStylewhite, Stigmawhite,
fundesulcato, stylo perlongitudinemin suLco involuto; 4-5 mm long,bilebed.
basalibiis cup shaped, ciliate around mouth. Fruitunknown.
labellum
concavum;
anthera versatilis calcaribus
nentes;
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APG
M)1.58
B
A
([glgl)
[!l)
Ii
I2mm
cm
D
5mm
cm
Ii
Ii
cm
E
F
z1:
;
'1tl
N111
1l[t
:11ll
F
1
:i1t'ai
E
il'1
l
.d
/t t
+
icmI
]
Fl1I,
FiG.1.deochlanots
mullerensis,
A: Crosssection of ovary, B[ Epigynousgland.
C: Anther D: Calyx.
E:Bract,
F: Bracteole.
G: Flawer.
H: Partof fiowershowing corolla tube with a furrowenclosing a style,lateral
staminodes,
dersalcorolla lobe,anther, and labellum (from
spirit-fixedmaterial ofH/ Ohada et al. K767J).
Distrihtttion:
West and CentralKalimantan.
JP8 CK);
ibidem,CS Awmaek J99(K).
wrst Kalimantan:
Serawai,8 km northeast ofDesa Jelundung,
BatuLmiang
112"36'7.1"E),AC Chureh & U]drMlabyar
(O"30'57.9"S,
Etymology:The highlyviscous mucus from
glandson the bracts is pooledin the axils of the
17e2 eo).
imbricatebracts,The name of the new genus is
derived
from these features
slimy + ehlanurgMblecularpbylogenetic analyses
(MJarocovering,
using
I71Sand
mantle).
combinedclatasets
The ITS-1had a tota1 aligned lengthof 184 bp
Additionalspecimens examined,
INDONESIA.
sequences
ranged
from 173 to 217 bp)
CentralKalimantan:Sarnba, 1994-1995, Cuttingbleck (unaligned
ofPT Handiyani(Oa43'22.5"S,
112'50'37,S"E),JKJti,'vie with a mean
guanine-cytosine
(GC)content of
& A Ruskandi 5447 (BO);Near Muara Joloi (Oe02'S 53,59t6,
the 5,8S region had an aligned lengthof
114"06'E), es Awmack 109 (K),
ibidem, {:USAwmack
164 bp
158-164bp) and a GC content
(range,
of
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TZiXKANO & NAGAMASU:
FIG.2.A(ytrochlam.vs
murlerensis.
"6,xochlanrys,
gen.nov.
23
A/ Habitat.
B: Close-upof fiower.C: Infiorescence
(all
photosby H.Okada.; H. Okada
etal.
KZ
67J).
51.79t6,
and the ITS-2 had an
aligned
bp (range,
208-270bp)with
GC
a
lengthef226
content
In toxochtaop,s,
the ITS-1 sequence
vv'as
of58.1%.
The result ofBayesian
inference
ofphy(58,4%),
logeny(posterior
PP), howevegstrongprobahilities:
179 bp in
ly supported
lengthwith a GC content of52.5%, the 5.8S region
was 164 bp inlength
with a GC content of51,8%,
and the ITS-2 region was 229 bp in Iength
with a
GC content of 59,89r6,
Analysis of the ITS sequence data,excluding
the ambiguously
aligned regions by using 70 taxa,
resulted
in eight equally parsimonioustreesof 1508
steps (number
of parsimony-informative
characters,272; consistency
index [CI] O.404;retention index [RI] O,622).In the strict consensus
tree (Fig,
3), Myxochlamys
is found in the
Zingiberoideae clade, which
is sister to the
Scuphochlamysand Distichochlam.vs
clades, but
bootstrapsupport for this relationship
is low
=
=
and
=
4).
(Fig.
Congruence of the datapartitions
(chloroplast
nuclear) could not be rejected by the ILD testlp
(PP
1OO) this relationship
=
O,98),Analysis ofa
2227
characters
combined
fbr47 taxa
matrix
containing
in 1617 equal-
resulted
ly parsimonioustrees of 1711 steps (number
of
characters, 401; consistency
parsimeny-informative
=:
-
index[CI] O.527;retention index[RI] O.603).
The strictconsensus treeisshown in Fig.5,Both
MP and Bayesiananalyses (Fig.
6)yieldedtreesof
similar topology,but the Bayesian tree was more
resolved
inthe Zingiberoideaeclade. toxochtamys
issister te Sbcrphochlam.vs
and
(BS!PP74.31100),
Distichochlaiays
issister to these genera (BS!PP
72.9flOO)(Figs,
5, 6).
-
-
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APG
Vbl.S8
Gtobba talangensis
Camptandraparvuta
Caulokaempteriasaxicola
Hanith'a
aloifiora
PJffgophyilumyunnanensis
Curcumorpha longifiora
2
aoo.D7o.a
Boesenbeigia iongifiora
100,O
Curcumorphatongifiora7
59.899.9riOD.OBoesenbeigiapuicherrima
Boesenbeigiagelatinosa
Boesenbeigia rotunda
Caulokaempieriathailandica
Com"kaempferiatongipetiolata
Comukaempteria a"rentifiora
Zingibereliipticum
Zingibergramineum
Zingibersuiphureum
Zingiberjunceum
98.850.758.0
95.4
Zlngiberoideae
9B,9
58,4
Clade
58.7
82.4
67.699,492.9
88.2
Zingiberwrayii
ZingiberJongipeduncuiatum
Zingibercoloratum
MJorochlamys mullerensis
Distichochlan?ys
sp. Kress O1-6848
Distichochlamysciteaf
Distichochiamyscitrea2
Scaphochlamyskunstierif
Scaphochlamysbiloba7
Scaphochlamysbiioba2
Scaphochlamyskunstleri2
Scaphochlamysobcordata
Kaerr7pteria
elegans
Kaernpteriaparvifiora
Kaempferia rotunda
99.7
97.5
97.4
95.599.9
Kaempteria sp. kress98-6289
Cautieyagrac"is
Cautieyaspicata
Roscoea bhutanica
Roscoeaputpurea
Roscoea cautieoides
Roscoeahumeana
Hea)fchiumlongicomutum
Hedychium
viUosum
Pommereschea
iackneri
Rhynchanthusbeesianus
Smithatris
scrpreneanae
8g.2
too,o
too.o52.8
92.3
84.0
80,572.994.7
loe.o
Curcuma bicolor
Curcuma ecomata
StahJianthusinvotucratus
Curcuma harrnandii
Curcumaparvifiora
Curcuma thorelii
Curcuma roscoeana
Paracautleyabhath'
Curcuma rubescens
Hitcheniaglauca
Hemiorchisrhodorrtrachis
Gagnepainia
hannandii
loa.o93.a
55.793.78d,3
77.1
rioo.o
53.9loe.o
Gagnepainia thoretiana
AipiniablepharocaCyof
Elettaria
longituba
Atnomum yunnanense
Aipiniatonkinensis
Aipiniaeiegans
Renealrriiabattenbergiana
Eiettariopsisstenosiphon
Eiettariopsiskerbyi
Tarnijia
fiageliaris
SiphonochiJusaethiopicus
Siphonochilus
kitVcfi
Siphonochilusdecorus
FIG,3,Strict
consensus
of eight maximum
resulting from analy sis of the ITS
parsimonioustrees of the subfamily Zingiberoideae
sequence
data(length1508,consistency index O.404,and retentjon index = e.622).Numbers above branchesindicateboot=
strap x,alucs
=
(>50%).
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llAKANO & NAGAMASU:A(yxochlanrys,
25
gen.nov.
Gtobbatalangensis
Oamptandiaparvula
Cautokaempferiasaxicola
PyrgophyitumHaniffiayunnanensis
aJbifiera
Boesenbergiageiatinosa
Boesenbergiapulcherrima
too73
10D
Boesenbergiarongiflora
Curcumorphatongifioraf
'
Boesenberg
aoo99CurcumorphaIongiftora2
Zlngiberoideae
Claderioo
iarot"nda
ga
riooCa"lokaempferiathaitandica
Cornukaempferialongipetiolata
Cornukaempferiaaurantifiora
Zingibereliipticum
92
Zingibercoloratum
leO
Zingibertongipeduncuiatum
Zingiberwray ii
Zingibersuiphureum
Zingibergrarnineum
Zingiherl'unceum
5259
96
5360
nayxochiamysmullerensis
Distichochlamyssp.KressOf-6848
aoo
leo
ioo
DistichochtamyscitreaV
Distichochtamyscitrea2
99
100
Scaphochlamysbitobaf
Scaphochlamysbiloha2Scaphochtamyskunstferi7
rioo
100Scaphochtamyskunstleri2
63
Scaphochlamysobcordata
Kaempferiaelegans
Kaempferiaparvifiora
66
100
{ooaoo
Kaempferiarotunda
100
Kaempferia
kress98-6289
aooCautieyagracilis
Cautleyaspicata
100
Roscoeabhutanica
riootooRoscoeapurpurea
looRoscoeacautleoides
Hedychiumrongicotnututn
100Roscoeahurrieana
86
Hedyohiemvilros"m
100
beesianus
99RhynchanfhusPomrrierescheaJackneri
Curcumabicolor
CurcumaSmithatrissupraneanee
ecomata
leo
C"rcumaharmandii
100100
60
StahJianthusinvolucratus
Curcumapatvifiora
ioe
Curcumathoretii
Curcumaroscoeana
9898100Paracautieyabhatii
Curcumerubescens
Hemiorchisthodorrhachis
riDO
GagnepainiaharmandiiHitcheniaglauca
92Gagnepainiathoretiana
Alpiniablepharocatyx
Reneatmia battenbergiana
Elettariopsiskerbyi
rioo
Eiettariepsis
stenosiphon
87
100
Eiettarialongituba
100
leo
Amomumyunnanense
100
Arpiniatonkinensis
Arpiniaelegans
Tami'iafiageilaris100
99
Siphonochitusaethiopi
Siphonochiiuskirkii
Siphonochiiusdecorus
O.fiSubstitutionsiSite
Flci.
4. Bayesianconsensus
phylogram based on branch lengthobtained
than 5U%.
(PP)higher
posterior
probabilities
fromITS scqucnce
data.
Nurnbersabove branches
indicate
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Camptandtaparvuta
Cauiokaempferiasaxicota
RyngophWlumyunnanensis
Boesenbeigiarotunda
Boesenbeigia
puichern'ma
Curcumorpha tongifiora1
Curcumotphalongifiora
2
Cornukaempleriaautentifiora
Zingtoerellpticum
Zingtoergramineum
Zingibersuiphureum
Zingiber
umayii
Distichochlamyssp. Kress Ot-6848
IPlyxochlamysmutlerensis
Scaphochlamys biloba
f
Scaphochlamyskunstieri1
Kaempforiaparvifiora
Kaempferiarotunda
Kaempforia sp. kTess98-6289
Cautleya
gracilis
Cautteyaspicata
Roscoeacautleoides
Roscoeapurpurea
,
Hedychium longicomutum
Hedychiunivillosum
Pommereschea lackneri
Rhynchanthusbeesianus
Curcuma bicoior
Smithatrisstq)raneanae
Curcuma roscoeana
Hitcheniaglauca
Curcuma thorelii
Stahtianth"sinvoiucratus
Gagnepainia
thoreliana
Hemiorehisrhodorri]achis
Aipiniablepharocalyx
Aipiniaelegans
Amomum viitosum
Amomum yunnanense
Renea/mia battenbeigiana
Elettariopsiskembw'
Etettariopsisstenosiphon
lamijiafiagelfaris
Siphonochilusaethiopicus
Siphonochilusdecorus
Siphonochiluskirkii
FIG,5,Strict
consensus
and
es
1617 maximurn
Zingiberoideae
resultin.tr from analysis ofcembined
parsimonious trees ofgubfamily
ITS
= L711,consistency indcx= O.527,and retention index= O.603).Numbers above branclidata(tength
indicate
bootstrapvalues (>50%).
tnatK
rcgion
of
sequence
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& NAGAMASU:ta・xochlam.vs, gen.nov.
Arr]omum yunnanense
Amomum villosum
Aipiniaetegans
Aipiniablepharocat)o(
evettariopsisstenosiphon
Elettariopsiskembyi
Renealmiabattenbeigiana
Gagnepainia thoreffana
Stahlianthusinvolucratus
Curcuma thoreM
Hitcheniagiauca
Curcuma roscoeana
Smithatrissupraneanae
Ryrgophytlum
yunnanensis
Camptandrapantula
Rhynchanthusbeestanus
2ZlngiberaLdea
Heclychiumvi"osum
Hedychiumlongicornutum
Roscoeaputpurea
Cautleyaspicata
Kaempleriasp.
Kaempteria rotunda
Keempteriaparvifiora
ScaphochiamyskunstieriV
Scaphochlamysbilobaf
na)orochlamysmullerensis
Distichochtarnys
sp. Kress Of-6848
Zingiberwrayii
Zingibersuiphureum
Zjngibergremineum
Zingiherettipticum
Cornukaempferiaaurantifiora
Curcumorpha
tongifiora2
Curcumotpha iongifiora7
BoesenbeigiapulcheMma
Boesenbergia totunda
alhifioraCaulokaempteriasaxicola
tamijiafiageliaris
Siphonochiluskirfc"
Siphonochilusdecorus
Siphonochiiusaethiopicus
O.1SubstitutionsiSite
FJG,6. Bayesian consensus phylograrnbased on branch lcngthof combined
branchesindicatc
posterior
prebubilities
(PP)higherthan 50%,
ITS and
matK
region
sequence
data.Numbers above
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TABLE1 .Comparison of keycharacters betweentoxochlatays
and related genera
Arrangement Modeof
ofbracts
fiowering
Flowerslst bracteoleCorollatube Thecae
Anther Ovary
SolitaryOpen to base, withagroove
Basalspurs
Oppositebract; formed by the
keels2
innersurface
Versatile
IICucochtanozs
Spiral
Base-apex
mutlerensis
(indeteuninate)
Unilocular
with basal
placentation
Sbaphochlanrys Spiral
Base-apex
Usually Open to base, without such a With very
Adnate
in
Oppositebract, groove
short, free,
(indeterminate)
cincinni eften more or
basalspurs
lcsskeeled(2
(nospur in
keelsonly inS.
S.calctcola)
Unilocular
or
trilocularwith
basalplacentation
catcicoia)
Distichochtamp's
Distichous
Base-apex
In
Tubular;
keels2
cincinni
(indeterminate)
withoutsucha
Spursabsent Adnate
Sbaphochlanrys
are moderately
basedon theirspecies alone
Discussion
Trilocular
groove
supported
subclades
sub(St]ophochlamrys
MJtxochlanoLs
mullerensis
clade:
BSIPP -= 67.61100; Distichochlatayssub-
oideae
clade:
BS!PP
isplacedintheZingiberin the phylogeneticanalyses basedon theITS
-
95.4/100),and
was
tocrochlanrys
placedoutside of these genera. In the combined
was foundto be
data(BSfPP 58.71100firomITS data,89.6flOO data set phylogeny,iLdyxochlantys
from the combined data).
sister to the Scaphochlamys clade
In terms ofmorphology,
(BSfPP
74,3/100)and Distichochlamys was the sister of
A(lyxochlantys
mullanensis could also be placed inthe
tribeZingibereae,
subfamily
Zingiberoideae,inthe these two genera (BSfPP 72.9/100).The results
classification system ofZingiberaceae
by Kresset aL suggest thatthesethree generaare closely related,
Morphologically,A(YxochlaTays
issimilar to
by yirtue of the parallelorientation of the
(2002)
of itsleafyshoots, its
Scaphochtamys in that itlacksa distinct
well-develplaneof distichy
pseudostem,but the spiral arrangement
of bracts
oped latera1
staminodes,
and 1ahellum
not connate to
difl
At BO and K, most specimens of toxochlanrys
the filament.
The unilocular ovary with basal
pla- fers.
centation
israther rare in this tribe, however; a without fiowerswere identified
as Sbuphochianrys,
transition in the ovary type within
some
genera, probablybased on inflorescenceand stem characteristics.Thereare also significant morphological
e.g., Boesenbergia, Hbptochorema, and 5Zraphochlanrys, from trilocular
differences,
however,between 7VrlyxochlatTrys
and
with axile placentationto
unilocular
with freeor basalplacentation
hasbeen Scaphocklamys,For example, flowers are borne
reported
singly inA{lyxochlanrys
vs. bornein cincinni; corol& Nagamasu 2000),
(Sakai
latube with a grooveenveloping the style insidevs.
Our molecular phylogeneticanalyses suggest
the aMnity of Adyxochlaniys to Distichochlanrys Iackngsuch a groove;labellum
concave
vs. bilobed
and Scophochlanqys (Figs.
3-6).Adyxochlanu,s
is or entire, but never concave; anthers versatile vs.
sister to the Distichochlannysand Scophochlanrys
adnate, Distichochlanrys
isunique and distinctfrom
in that ithas tubular bracteoles
clades as shown
by theITS dataset analysis (Figs.?Lfyxochlanrys
and
3-4),although the relationships among
arranged
bracts (Newman 1995).
thesethree distichously
Furthermore, DistichochlaTayshas adnate anthers
subciades
are unresolved
in both the MP and
lackingspurs, producesflowersin cineinni, and
Bayesiantrees.Further,bothDistichochlanop?s
and
and
the cembined
ITS and
matiZf region
sequence
=
=
=
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29
ndyxochlanv,s,
gen.nov.
7Llyxochlainys Doyle, J,J,& J,L, Doyle.1987.A rapid DNA isolation
has a flat,
bilobedlabellum,
while
of freshleaftissue,
procedure forsmall quantities
hasversatile anthers with longspurs, producesa sinPhytochem.
Bull.
19:
11-15.
and hasa concave, entire labelgleflowerperbract,
Fanis, J, S,,M. Kallerajo,
A. G. Kluge & C. Bult.1995,
lum, Comparison of key characters
between
Constructing
a significance test fbrincongruence,
Myxochtamys and the other genera is shown in
Syst Biol.44:570-572.
Table 1,Among these characters,
the versatile
an
Felsenstein,
J.1985.Confidence1imits
on phylogenies:
approach
using the bootstrap,Evolution 39: 783anthers are rather rare in theZingiberaceaeand are
791,
found only in five generabesidesMyxochtanv,s:
Ha]1,T, A. 1999. BioEdit: a user-frielldly biological
Camptandra, Cautleya,Curcuma, f]beracautlaya,
sequence
alignment
editor and analysis program fbr
and Roseoea(Larsen
et al. 1998),
The groovewithWindows 95f98fNT.Nucl. Acids Symp, Ser.41:
inthecorolla tube isalso uncommon
and so faris
95-98.
known only in themonotypic Slddiochilusand a few Huelsenbeck,J. P. & F. Ronquist.2001. MRBAYES:
1980).
Bayesian inferenceof phylogeny.Bioinformatics
species of Heclychium
and Zngiber (Smith
17:754-755,
Myxochlamys might be placed in ScaphoKress,
W. J,,L. M. Prince& K, J.Williams. 2002. The
ehlantys as a peculiarspecies, but we hesitated
to
of the gingers
phylegeny and a new classification
expand
the circumscription
of the latter
genus
evidence
from molecular data.
(Zingiberaceae):
becauseofpoor support from molecular phylogeAmer. J,Bot, 89: 1682-1696,
netic
analyses
and
such
significant
morphological
A.-Z.Liu,M. F.Newman &
,
and
differences,
Furtherstudy of toxochlamys
related genera
isrequired beforefirm conclusions on
itsplacementcan be reached,
gratefu1 to Pro£ HiroshiOkada, Direetorof
Betanical Gardens, Facultyof Science,Osaka City
We
are
mo]ecular
phylegeny
ofAipinia
Li, 2005. [lhe
Q,-J.
a
(Zingiberaceae):
Amer, J,
po)yphyletic
genusof gingers.
Bot.92: 167-178,
Larsen,K.,J.M. Loek,H. Maas & P.J.M. Maas. 1998.
Kubitzki, K. (ed.),
The families
Zingiberaceae,
1}i:
and generaofvascular plants.4 : 474-495.SpringerVerlag,Berlin.
. & M, F, Newman.
2001. A new species of
Distichochlanrysfrom Vietnam and some observacomplex
and
Universityforinviting
us to panicipate
in the Muller
Range expedition. We also wish to express our cordial
LIPI, Scientific
Academy of
thanks to Dr.Dedy Darnaedi,
tions on genericlimits in Hedychieae (ZingiIndonesia, lndenesia, and Dr. Irawati,
Directorof the
Nat. Hist. Bull. Siam Soc, 49: 77-80.
beraceae),
Bogor Botanic Garden,LIPI, Indonesia,
fbrtheir kind
A,
Liston, & J.W. Kadereit 1995.ChloroplastDNA
assistance
duringthe fieldtripto CcntralKalimantan,
and long distance
disevidence fbrintregression
and Ms, Titien
Praptosoewiryo
who supported our fieldin
the
desert
annual
Senecio
persal
.flavus
wotk. Thanks are also due to the directors
and curators of
Plant,Syst.Evol.197:33-41.
(Asteraceae).
BO, E, and K fbrailowing us to examine herbarium
Manos, R S, & K. P. Steele.
1997,Phylogcnetic
analyses
specimens
in their care. Dr, Resalind Searle provided
of
Hamamelididae bascdon plastid
critical comments
on an earli ¢ r vcrsion of the manusequence data.
Amer. J.Bot, 84:1407-1419.
script. Thisgtudy was partlysupported by research
grants
McNeill,J,,F. R, Barrie,H. M. Burdet, Vl Demoulin,D,
from the Mitsubishi
Foundation
in the NaturalSciences
L, Hawksworth, K. Marhold, D. H. Nicolson,J.
and the Japan Societyfbr the Promotion of Science
Prado, P. C. Silva,J.E. Skog,J.H, Wiersema & N.
TropicalBio-resources
ResearchFund.
(JSPS)
J, Turland (eds,),
Inteniational
code of botanical
nomenclature
Code), A, R, G. Gantner
(Vienna
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TAKANO
Appendix 1.Genbank accesien numbers
and
& NAGAMASU:
youcher
specimens
A45ixochlamys,
gen.nov.
used
31
in this study.
Species
ITS
Aipiniablq;?harocalyx
K. Sehum,
AF47S709AF47S713AYT423S6AY769828AY352e12AY424744AY42474SAF47S725AF47S727AF47g730AY47873
AF478809AF478813
Kresset aJ, 2002
matK
Aipiniaelegans K.Schum.
AipiniatonkinensisGagncp.
Referenees,iVouchcr
Kresset aL 2e02
Kresset al. 200S
AmomuntvillosumLour.
AF47S824AY352042
Hanis et
aL
200S(1TS),Kresset
Amomum yunnanenseS, Q,Tong
Xiaet aL 2004
BoesenbergiagelatinosaK.Larsen
Ngainriabsakul
et at. 2003
BoesenbergialongijZoraKuntze
Ngamrtabsaku]et al. 2003
AF47Sg2SAF47S826AF47S83eAF47Sg31
Kresset al,2002
Boesenbenygi'apulchem'imaKuntze
Boesenbergiarotunda(L,)Mansfi
Kresset al. 2002
Kressetai, 2002
CanrptandraparvutaRicl1,
CautohaenjleriasexicoJaK.Larscn
Kresset
CauiohaemnferiathaigandicaK.Larsen
Ngamriabsakal
et aL 2003
Cautieyagraeilis(Sm.)Dandy
al,
2002
AF47S833AE478834AE47883S
Kresset aL 2002
CautieyaspicataBaker
Krcssetal.2002
Krcsset al. 2002
Coi'nuimeiqntleriaaut'antijioraJ,Mood&K,Larsen
Cornukaemrv2irialongipetjolataJ.Mood&K.Larsen
Curcuma bicolor
J.Mood & K. Larsen
Ngamriabsaku1et
AF47S837
al.
2003
Ki'ess
et al.2002
Curcumaecomata Craib
CuffcumaharmandiiGagnep,
Ngamriabsakulet al, 2003
CurcumaparvgeoraWall.
tv'garuriabsaku1
et at. 2003
Kressetal. 2002 CITS)
/Cuoet
Ngamriabsaku]et
CurcumaroscoeanaWal1.
CurcumarubeseensRoxb.
CurctfmathoreiiiGagnep,
AB047741
Di,stichochlantys
citrea M. F. Newman 2
Disiichochlanv,ssp.KressO1-6g4g
al,
unpublished
(matK)
Kressetal,2002
Kressetai.2002
Kresset
al,
2002
NgamriabsakLiletaL2003
AF47S844
Eiettarialongituba(rudl.)Holtturn
Eiettar'iopsis
kerbyiR, M. Sm.
(K.Schurn.)B. L. Burtt&
R. M, Srri,
Krcsset
Sakaiet
aJ.
2002
al. unpublished
AF47SS45AF47gS47
Pedersen2004 (ITS)
,i
Kresset
Kresset al.2e02
Gagnepainia
iJarmandii
K. Schum.
al.
2002 (matK)
Williains
et al, 2004
GagnepainiathorelianaK.Schum.
AF47gS51
Globba taiangensis A, Takano et H, Okada
filrnmpaatbijleraK.Larsen&J,Mood
Hbd.vchiumlongicornuium
Griff ex Baker
Kressetal, 2e02
Williaills
et aL 2004
AF47S855AF47SS60AF47SS61AY341090AF47Sg64
Kressetai.2e02
Kresset aL 2002
Hed),chiumvjllos"mWall,
Kresset al, 2002
Williamsetal.2004
K, Schum,
Kresset
KlteiptriaelegansWatl,
Kkeemnferiaparvt"of'aWal].
aL
2002
Ngamriabsaku1
etal. 2003
AB232052AF47gg68AF47SS69AB269791
Searle&HeddersonZOOO
KlrtenrpteriarotundaL.
Kaemnferiasp,kresb'9S-62SP
Alyxochtano,s
mullerensis
A,Takano&
2003
AF47S841AF47SS42AF47S843
Krcsset al. 2002
Verma1
Curcumorpha longgeora(WalL)
A, S, Rao & D. M. Vermu 2
Disiichochiantys
citrea M, F.Ne-man 1
llbmiorchis
fhoderrhachib'
th't:henia
glau:aWa[t.
aL
Ngamriabsakul
et al, ?O03
Curcumotpha longijlora
A. S. Rao & D. M,
(Wa]l.)
Eiettariopsis
stenostPhon
2002 (matK)
aL
KressetaL 2e02
Kressetai.2002
Nagam,
Thisstudy: H, Okada et al.KIL671
NgnmiTiabsakn1
et aL 2003
Paracazattq,a
hAatii
R, M. Sm.
PommereschealackneriWittm,
AF478877AF47SH78AF47S880AF47S88S
Kresset al, 2002
P.Frgephytlune
T. L. Wu & Z. Y. Chen
)ninnanensis
1{eneainvia
battenbejgiana
Cummins ex Baker
RkynckanthusbeesianusW.W.Srn,
RoscoeabhutttnicaC.Ngamriabsaku1
RoseoeacautteoidesGagnep.
Roh'coeahumeanaBalf.f,
& W. W. Sm
Ros'coeapurptfreaSm.
thaphochtantvsbiloba(rudl,)HoltLam
Kressetal. 2002
Kresset
AF478887
2002
al.
20e3
Kresset at,2002
Wood et al. 2000
AF47gg88AY4788S9
Kresset al.2002
Krcsset al,2002
Scaphochlam.vsbiloba(Ridl.)Holttum2
Sbaphochlam.vslaunstleri(Bakei}Holttum
aJ,
Krcsset aL 2002
Ngamriabsaku]et
Wood
AY478890
et aL
2000
Kresset al.2002
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APG
Nio1.58
Appendix 1.continued,
Species
ITS
SbcQvhochtafayskunstteri(Baker)Holttum
AJ3SS2S7
AJ3gS2S6
Scaphochlan-'sobcordataP.Sirirugsu&K.Larsen
Siphonochilusaethiopicus(Schweinf,)B,L,Burtt
matK
ReferenncesiVoucher
Searle&1・Ledderson2000
Searle&Hedderson2000
M478893AF47g894AF478S95AF478896AF478897AF47gB9g
Kresset aL 2002
Kress et aL 2002
Sij]honochilusdecoruh'([lruten)Lock
AF478792
AF478793
Siphonochilusldrldi{Hook,)B,L.Burtt
AF478794
Kresset
Sinithatris
supraneanae W. J.Kress&K, Larsen
involttcratus
ex Baker)R. M. Srn.
Stahiiantkus
(King
AF478795
AF478796
Kresset al.2002
7lamijiafugellarisS.Sakai&Nagarn.
AF478797
Z'ngibercoJorainmN.E,Br,
AF414498
Kresset aL 2002
Pedemsen20pa CrTS)
Znigber elij)tieum
(S,
Q.Tong &
Y. M. Xia)S,Q,Tong & Y. M. Xia AF478799
ZingibergramineumNoronha
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