E DIN B . J . B O T. 57 (2): 261–270 (2000)
261
PHY LOGE NY O F H ED YCH I UM A ND R EL ATED
G EN ER A (ZI NG IBE RA CEA E ) BA SED O N IT S
SEQUENCE DATA
T . H . W OOD *, W. M . W HIT T EN † & N. H. WIL LI AMS‡
The phylogeny of Hedychium J. Koenig was estimated using sequence data of internal
transcribed spacer regions 1 and 2 (ITS1, ITS2) and 5.8S nuclear ribosomal DNA.
Sequences were determined for 29 taxa, one interspecific hybrid of Hedychium and one
species in each of 16 other genera of Zingiberaceae representing tribes Hedychieae,
Globbeae, Zingibereae and Alpinieae. Cladistic analysis of these data strongly supports
the monophyly of Hedychium, but relationships to other genera are poorly supported.
Within Hedychium, four major clades are moderately supported. These clades are also
distinguishable on the basis of number of flowers per bract and distribution.
Stahlianthus, Curcuma, and Hitchenia also form a strongly supported clade. Based on
this limited sample, the currently defined tribes of Zingiberoideae are not
monophyletic. The Asiatic genera form a monophyletic group within this broadly
defined Hedychieae. The taxonomy and biogeography of Hedychium are reviewed.
Keywords. Biogeography, ginger, Hedychium, ITS, phylogeny, Zingiberaceae.
INT ROD UCT ION
Within the Zingiberaceae four tribes are currently accepted (Smith, 1981): Globbeae
(bow-shaped filament with unilocular ovary with parietal placentation; four genera);
Zingibereae (pointed anther crest surrounding the style and staminodal lobes adnate
to the labellum; one genus, Zingiber); Hedychieae (plane of leaf distichy parallel to
the rhizome, lateral staminodes petaloid and free from the lip except in Siphonochilus
and Curcumorpha, ovary trilocular with axile placentation or unilocular with basal
or free columnar placentation; 21 genera); and Alpinieae (distichy of leaves perpendicular to the rhizome, lateral staminodes reduced to teeth or swellings or absent,
ovary trilocular with axile placentation; 22 genera). Alpinieae have no capacity to
shed the stems or inflorescences by abscission. The other three tribes readily shed
their stems and form a corky abscission layer on the rhizome in response to old age,
photoperiod, soil temperature or drought. The position of Hedychium within tribe
Hedychieae is uncertain.
The tribe Alpinieae is pantropical, ranging from New Guinea and Fiji in the East
through Asia, Africa, and Central and South America ( Wood, 1991; Wu, 1994). The
other three tribes occur mainly in southern Asia with sparse representation in Oceania
(with no truly indigenous species east of the Moluccas). The only exception is
* Department of Environmental Horticulture, University of Florida, Gainesville, FL 32611, USA.
† Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA.
‡ Florida Museum of Natural History, University of Florida, Gainesville, FL 32611, USA.
262
T . H . W OO D E T AL .
Siphonochilus which is endemic to Africa. All fossil Zingiberaceae, some of which
date from the late Cretaceous, have been interpreted as having affinities with
Alpinieae ( Friedrich & Koch, 1970; Hickey & Peterson, 1978).
The tribe Alpinieae are all tetraploids with 2n=44, 48, 96. Globbeae are diploids,
tetraploid, hexaploids, and octaploids with basic numbers x=8, 10, 12. Zingibereae
are all 2n=22. Hedychieae are mostly diploid but with x=9, 10, 11, 12, 14, 17
(Hedychium, Cautleya), 21, 25 (Chen, 1987, 1989).
The taxonomy of the horticulturally important genus Hedychium has been controversial since the middle of the nineteenth century. At least 115 names have been
published in the genus; we currently estimate that about 65 of these are biologically
valid species. The species are concentrated in northeast India (24 spp.), Burma (11),
and in China [in Yunnan province (18), Guangxi province (8), Guangdong and
Hainan provinces (2)], northern Vietnam (10), Nepal (12), northern Thailand (14),
peninsular Malaysia (7), southern India (4), Borneo (4), the Philippines (2), Java
(3), and Moluccas (1).
The terrestrial species in the circum-himalayan region grow in cool, wet mountains
up to 2400m, whereas the Malesian species are mostly epiphytes. Species delimitation
has varied among authors; for example some divide H. coronarium into 9 species,
and H. coccineum into 7 species (Roscoe, 1828; Turrill, 1914). All authors agree that
the genus is monophyletic. It is characterized by long, linear or lanceolate lateral
staminodes, persistent, coriaceous bracts, and a long, exserted stamen with a lower
dorsifixed anther. The labellum is showy and usually emarginate or cleft into two
lobes.
Wallich (1853) circumscribed the following subgeneric divisions: Coronariae (more
or less tightly imbricate spikes); Spicatae (elongated spikes with distant, spreading
bracts); Siphonium (one species, H. scaposa, a slightly crested anther and a stemless
habit similar to the genus Kaempferia to which it has long since been transferred );
and Brachychilum (H. horsfieldii, a cleistogamous plant almost lacking a labellum
with two wide lateral staminodes, formerly a segregate but recently placed within
Hedychium (Newman, 1990). Horaninow (1862) divided Hedychium into three
groups: Gandasulium (stamen shorter than or equal to the length of the labellum),
Macrostemium (stamen much longer than the labellum), and Brachychilum (one
species, lacking a significant labellum). He also classified four Indonesian species as
incertae sedis. Schumann (1904) redefined Gandasulium to include taxa with a dense,
short and wide, ellipsoid, ovoid, rarely long cylindrical inflorescence; with bracts
flat, densely imbricate, rarely arched; and with rachis always hidden. He defined his
other subgenus, Euosmianthus, to include species with less dense, with longer than
wide inflorescence; bracts never densely imbricate, commonly patent, or divergent,
or distant from each other, clasping the flowers; and with rachis not hidden. He also
maintained a separate genus for Brachychilus, comprising a Moluccan species and
H. horsfieldii. It is the aim of this paper to examine the validity of these classification
schemes in light of modern molecular phylogeny.
The first author is writing a monograph of Hedychium. Preliminary phenetic analy-
PH YL O GE NY O F HE DY CHIU M
263
sis of 15 species ( Wood, 1996) using 11 inflorescence characters showed some support
for grouping of the species into imbricate and tubular bracted groups. Later analysis
( Wood, unpublished data) using 110 specimens of 67 species did not indicate wellsupported clustering of the two bract types in factor analysis; however, discriminant
analysis correctly scored bract types in 90% of the observations and five of the eleven
misclassifications involved Malesian species. The only published molecular studies
of Zingiberaceae use rbcL sequences (Clark et al., 1993; Kress, 1995). In these analyses Costaceae are unresolved among the other five families of Zingiberales and only
the four tribes of the Zingiberaceae form a single clade. When morphological characters are added to the rbcL data the Alpineae becomes the outgroup to the other
three tribes.
Wood (1991) hypothesized that the Costaceae and Alpinieae are the earliest groups
in the Zingiberaceae and originated in Western Gondwanaland before the effective
separation of South America and Africa, although fossil evidence from North
America and Europe run counter to this hypothesis. These two groups were rafted
on the Indian subcontinent from Africa to Asia. Sometime when the subcontinent
was in the middle latitudes, when the paleoclimate was fairly dry, the progenitors
of the other three tribes of the Zingiberaceae evolved in response to climate, and the
ancestors of the African genus Siphonochilus dispersed to eastern Africa. Upon the
collision of India with Asia, the uplift of the Himalayas provided many isolated and
seasonally favorable habitats that prompted a massive radiation of genera of the
Hedychieae, Globbeae, and Zingibereae in upland areas while the Alpinieae flourished
in the lowland tropics of Asia and Oceania.
MAT ERI ALS AN D ME TH ODS
Leaf samples of 29 species and cultivars of Hedychium plus representative species
from 16 genera in the tribes Hedychieae, Globbeae, and Zingibereae, and one member
of the tribe Alpinieae were obtained from material cultivated by the first author. The
Alpinia was selected as the outgroup based on Kress (1995), fossil evidence, and the
biogeographic evidence cited above. Also, one specimen that was thought to be an
interspecific hybrid (Schilling, 1982) and a wide interspecific hybrid created by hand
pollination were included in the analysis in order to evaluate our ability to detect
natural interspecific hybrids. A list of taxa examined and voucher numbers is presented in Table 1. Vouchers are deposited in the University of Florida Herbarium
( FLAS).
Fresh or dried tissue was ground and extracted by the modified CTAB method
(Doyle & Doyle, 1990). The ITS1 and ITS2 regions along with the intervening 5.8S
nrDNA region were amplified using PCR with primers 5 and 4 of Baldwin (1992).
The amplified products were cleaned on QiagenA columns according to manufacturer’s instructions. Dye terminator cycle sequencing reactions were performed using
Applied Biosystems reagents and protocols. AutoAssembler software (Applied
Biosystems) was used to assemble the complementary strands and edit nucleotide
264
T . H . W OO D E T AL .
TA BL E 1. Taxa sampled
Species
Group
Voucher
Hedychium acuminatum Ker Gawl.
H. borneense R.M. Sm.
H. bousigonianum Gagnep.
H. carneum sensu Y.Y. Qian
H. coccineum Ker Gawl.
H. coronarium König
H. cylindricum Ridl.
H. densiflorum Wall.
H. densiflorum ‘Stephen’
H. elwesii Baker
H. ellipticum Ker Gawl.
H. flavescens Roscoe
H. gardnerianum Roscoe
H. glabrum S.Q. Tong
H. gracile Roxb.
H. gracillimum A.S. Rao & Verma
H. greenii W.W. Sm.
H. hasseltii Blume
H. horsfieldii Wall.
H. longicornutum Baker
H. muluense R.M. Sm.
H. maximum Roscoe
H. puerense Y.Y. Qian
H. spicatum Ker Gawl.
H. stenopetalum Lodd.
H. tenuiflorum K. Schum.
H. thyrsiforme Ker Gawl.
H. urophyllum Lodd.
H. yunnanense Gagnep.
H. hasseltii×gardnerianum
Alpinia intermedia Gagnep.
Boesenbergia aurantiaca R.M. Sm.
Cautleya robusta Baker
Cornukaempferia aurantiflora J. Mood & K. Larsen
Curcuma petiolata Roxb.
Gagnepainia godefroyi K. Schum.
Globba pendula Roxb.
Haniffia cyanescens Holttum
Hitchenia glauca Wall.
Kaempferia roscoeana Wall.
Pommereschea lackneri Wittm.
Roscoea humeana Balf. & W.W. Smith
Rhynchanthus beesianus W.W. Smith
Scaphochlamys biloba Holttum
Siphonochilus kirkii B.L. Burtt
Stahlianthus involucratus Craib
Zingiber corallinum Hance
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Alpinieae
Hedychieae
Hedychieae
Hedychieae?
Hedychieae
Globbeae
Globbeae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Hedychieae
Zingibereae
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
T.
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
Wood
14
18
11
35
6
15
13
24
47
10
29
37
8
16
35
36
26
23
2
9
12
27
30
38
28
7
25
40
17
49
1
32
51
19
50
21
31
38
43
20
45
46
32
33
41
44
34
PH YL O GE NY O F HE DY CHIU M
265
sequences. Sequences were easily aligned manually. The sequences are deposited in
GenBank (accessions AF202374-AF202420) and the aligned matrix is available from
the first author. PAUP* 4.0 b2 (Swofford, 1999) was used for parsimony analysis.
Initial analyses consisted of 1000 replicates of random taxon addition using SPR
and MULTREES, saving only three trees per replicate. These trees were then
swapped to completion, or until 10,000 trees were saved. The data set was then
subjected to three rounds of successive weighting (reweighted on rescaled consistency
index; 1000 replicates, saving 5 trees per replicate) to decrease the effects of highly
homoplasious sites. Lledó et al. (1998) gave convincing reasons for using successive
weighting. Successive weighting reduces the effects of highly homoplasious sites, and
thus emphasises sites that are more consistent.
In addition to maximum parsimony analyses, we evaluated support of the clades
by: 1, bootstrap analyses (Felsenstein, 1985) to obtain bootstrap support for nodes
using both equally weighted and successively weighted trees with 1000 replicates of
bootstrapping using SPR swapping, MULTREES on, holding 10 trees/replicate; 2,
by use of Bremer support (Bremer, 1988, 1994) to obtain branch support for the
equally weighted trees using the program Autodecay, version 4.0 ( Eriksson, 1998)
and PAUP* 4.0b2 (Swofford, 1999); and 3, by the reliability percentages obtained
by the quartet puzzeling method of maximum likelihood (ML) (Strimmer and von
Haeseler, 1996) as available in PAUP* 4.0b2. The ML parameters were the H-K-Y
model, ti/tv=2, with other parameters set to default. Bremer support trees and ML
trees were drawn in the TREEVIEW program (Page, 1996).
RE SU LTS AN D DISCU SSION
The aligned matrix is 733 base pairs (bp) long, and consists of the 3∞ end of 18S
(30bp), ITS1 (235bp), the 5.8S region (165bp), ITS2 (238bp), and 16bp of the 26S
region. Twenty-four bases were excluded due to ambiguous alignment. Of the 709
included sites, 330 were variable and 169 parsimony informative.
The initial unweighted analyses yielded 10,000+ trees with a length of 726 (CI=
0.618, RI=0.644). After three rounds of successive weighting (100 replicates, saving
5 trees per replicate), the final analysis produced 500 trees of length 315.56 (CI=
0.875, RI=0.838). The Fitch length of the successively weighted trees (equal weights
reapplied ) was 728 (CI=0.617, RI=0.641), two steps longer than the shortest
unweighted trees. The CI for unweighted trees with uninformative characters
excluded was 0.485, and the CI for weighted trees with uninformative characters
excluded was 0.690. Fig. 1 shows a randomly-chosen reweighted tree with branch
lengths above the lines, bootstrap support values (if 70%) directly under lines, and
decay values (given as d=) below the bootstrap values. Decay values are only given
for major clades if 1.
The monophyly of Hedychium is highly supported (bootstrap=100%). The tree
in Fig. 1 shows four moderately supported clades within Hedychium and illustrates
poor resolution within these clades due to low sequence divergence. The four clades
266
T . H . W OO D E T AL .
F IG . 1 . One of 500 reweighted equally most parsimonious trees with branch lengths given
above the lines and the bootstrap support values directly below the lines, with weighted
bootstrap values followed by unweighted bootstrap values. Weighted boostrap values are not
given for clades with values <70%. Decay values are below the bootstrap values and indicated
by d=(value). Additional statistics are given in the text. Genera not in the Hedychieae are
indicated by (Z, Zingibereae), (G, Globbeae), and (A, Alpinieae).
PH YL O GE NY O F HE DY CHIU M
267
form an unresolved polytomy in the bootstrap analysis. Clade I plants occur only
in southern Vietnam, the Malay Peninsula, and Oceania. They are short, generally
epiphytes or calciphiles, with a short day or day neutral photoperiod, slender inflorescences, one or two flowers per bract (three in H. bousigonianum), and flowers much
exserted from the bracts. Clade II is sister to Clade I in the successively weighted
strict consensus tree, and these two clades have weak decay support (d=1). Clade
III, represented here by only H. acuminatum, is sister to clades II and I in the strict
consensus tree (not shown). Clade III and Clade II are high altitude Himalayan
species that have only one flower per bract and a strict dormancy requirement. Clade
IV species have a wider circum-himalayan distribution at lower altitudes than Clades
II+III. A comparison of collection information from 23 herbarium specimens from
Clade II and 43 specimens from Clade IV showed a mean altitude for Clade II of
1783m and 1260m for Clade IV. This difference is highly significant (a<0.0005)
using a students t-test. These tall plants do not normally go dormant in the winter
and have three or more flowers per bract.
Because artificial interspecific hybrids of Hedychium are easily created ( Wood,
personal observation), natural hybridization is a potential source of taxonomic confusion in Hedychium. The ITS sequence of the artificial hybrid (H. gardnerianum ×
hasseltii ) was intermediate between that of the two parents. The parents differed by
17 bases; at each of these positions, the hybrid sequence displayed polymorphic
states, indicating ITS copies from each parent. These positions were scored using
ambiguity codes. When the hybrid sequence was included in the PAUP analyses (not
shown), the hybrid was sister (on a zero-length branch) to one or the other parent,
depending upon the addition-sequence replicate. In contrast, H. densiflorum and the
aberrant variety named ‘Stephen’ possess nearly identical sequences and are sister
taxa in the analyses with high bootstrap support. The sequence of cultivar ‘Stephen’
revealed no ambiguous sites. The similarity of these two sequences does not support
the hypothesis that ‘Stephen’ is a cultivar of recent hybrid origin (Schilling, 1982)
and suggests that it is merely an aberrant form of H. densiflorum. These examples
show that ITS sequences may reveal recent interspecific hybridization that might
otherwise confound phylogenetic analyses. Because ITS regions are known to
undergo rapid concerted evolution (Baldwin, 1992), species of ancient hybrid origin
may be difficult to detect without additional lines of evidence.
On a higher taxonomic level, a clade consisting of Cautleya, Pommereschea,
Rhynchanthus, and Roscoea is weakly supported (61% bootstrap), but not as the
sister group to Hedychium as we had expected on the basis of morphology. The
clade of Pommereschea and Rhynchanthus has a bootstrap support of 94% and a
decay value of d=5, with both values indicating very strong support for this clade.
This supports the idea that these two genera are in the tribe Hedychieae not the
Alpiniae as Schumann indicated based on the lack of lateral staminodes. This classification has been supported by Smith (1980) and by Z.Y. Chen (personal communication). They both lack petaloid staminodes, making them quite different from
Hedychium. Roscoea is sister to Cautleya with 92% bootstrap support and a decay
268
T . H . W OO D E T AL .
value of d=7. These are high altitude Himalayan taxa that bear single flowers in
each bract. The flowers have oblong, petaloid lateral staminodes and a bifid labellum
like Hedychium. The four genera together form a weakly supported clade.
Other related clades, while not reflecting on the phylogeny of Hedychium, do shed
light on the evolution of the Zingiberaceae. A clade consisting of Hitchenia, Curcuma,
and Stahlianthus has a high bootstrap support of 89% and very strong decay support
of d=7. The separation of Hitchenia from Curcuma on the basis of exserted flowers
and non-versatile anthers has never seemed adequate, but Stahlianthus (single bell
shaped bract) perhaps should be considered as a Curcuma with two adnate bracts.
The position of Zingiber makes the tribe Hedychieae polyphyletic. In spite of
describing the genus Cornukaempferia as having an ‘anther crest (that) shows a
striking resemblance with the anther appendage characterizing Zingiber’, Mood and
Larsen (1997) placed it close to Kaempferia on the basis of vegetative habit. The
ITS data indicate that Cornukaempferia is sister to Zingiber (92% bootstrap support)
and might not be generically distinct. The placement of Gagnepainia and Globba
make tribe Globbeae paraphyletic. It is notable that the Asian species of Globbeae,
Zingibereae, and Hedychieae form a monophyletic group with a bootstrap support
of 99% and decay support of d=3. The fact that the African genus Siphonochilus
appears as the sister group to the rest lends credence to the theory of an origin of
these tribes on the Indian subcontinent 40–50 million years ago.
The results of the quartet puzzling analysis also provides strong support for the
monophyly of Hedychium, which has a reliability percentage of 100. Within
Hedychium, clade II has a reliability of 94%, clade IV 99%, and clade I 98%. The
other clades with high puzzle support are Boesenbergia clade (83%), Pommereschea/
Rhynchanthus (92%), Stahlianthus/Curcuma/Hitchenia (98%), Curcuma/Hitchenia
(98%), and the large clade of Asian species (86%).
In conclusion, sequence data show that the genus Hedychium is monophyletic and
includes the genus Brachychilum. We propose these clades which may be described
later as subgenera: 1, Clade IV (three to five flowers per bract) and 2, Clade II (one
flower per bract), both of which have a natural circum-himalayan distribution; and
3, Clade I (one or two flowers per bract) that occur only in the Malay Peninsula,
Philippines, Borneo, Sumatra, Java, Sulawesi, and the Moluccas. The position of
Clade III (H. acuminatum) is uncertain but probably includes H. venustum (not
available for this study) on the basis of morphology. This molecular phylogeny runs
counter to Schumann’s classification because species in three of these clades occur
in each of his subgenera. This analysis also emphasises that the most important
factor in the evolution of this genus is geographic and ecological isolation. Because
tribes Globbeae and Zingibereae make tribe Hedychieae paraphyletic we recommend
that only tribe Zingibereae be retained. The position of the African genus
Siphonochilus is uncertain and must be evaluated along with other genera in the tribe
Alpinieae and Costaceae, preferably including African species, to determine its placement. This data set suggests that tribal concepts need to be re-evaluated in the
Zingiberaceae. Expanded data sets, such as the conserved matK, need to be used
PH YL O GE NY O F HE DY CHIU M
269
before the existing taxonomy is radically altered. The sister group of Hedychium is
uncertain based on this sample.
ACKN OWL E DG EM EN TS
We thank Bijan Dehgan for reallocating his resources for final sequencing. We are
also grateful to all of the institutions and individuals who provided living and leaf
material. We acknowledge the help of Li Qingjun of the Xishuangbanna Botanic
Garden, John Kress of the Smithsonian Institution, and D. Wankhar of the
Meghalaya Museum of Entomology for help in field research. We also thank Graham
Mallette, The Heliconia Society of South Florida, Jesse Durko, Glen Stokes, Karen
Taylor, Trish Frank, Donald Angus, Russel Adams, and Ken and Lyn Spencer-Mills
for financial help in collecting and Wendy Zomlefer and Walter Judd for editorial
comments. Portions of this work used equipment and materials funded by National
Science Foundation grant DEB 9509071 to WMW and NSF grant DEB 9815821 to
NHW. This is Florida Agricultural Experiment Station journal series number
R-07246.
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Received 5 August 1999; accepted with revision 15 February 2000