Plant Systematics and Evolution (2018) 304:1165–1180

https://doi.org/10.1007/s00606-018-1539-7

ORIGINAL ARTICLE

Taxonomic studies of Amomum (Zingiberaceae) in China II: transfer

of Hornstedtia tibetica to Amomum and supplementary description
of H. hainanensis
Xing‑Er Ye1,2 · Lin Bai3 · Yu‑Shi Ye4 · Nian‑He Xia1 · Jana Leong‑Škorničková5 

Received: 16 January 2018 / Accepted: 22 August 2018 / Published online: 27 September 2018
© Springer-Verlag GmbH Austria, part of Springer Nature 2018

Abstract
Hornstedtia tibetica was originally described only from fruiting material. On the basis of morphological study of flowering
material originating at the type locality, the species is a member of Amomum. This was also confirmed by palynological
evidence and a molecular analysis based on ITS region. A new combination Amomum tibeticum is therefore established
here, together with a detailed description and colour plate of this species. As previously suspected from the similarity of
the original description and type, Hornstedtia arunachalensis is conspecific with A. tibeticum and is therefore synonymised
here. The only remaining species of Hornstedtia in China is therefore H. hainanensis, which was also described only from
fruiting material. To aid unambiguous understanding of this taxon, we provide a detailed description and colour plate based
on recent collections of flowering material near the type locality in Hainan Province, China.

Keywords  Hainan · Lectotype · Synonym · Xizang

Introduction

A general introduction to Amomum Roxb., focusing on the

Handling editor: Karol Marhold.
* Nian‑He Xia
nhxia@scib.ac.cn
* Jana Leong‑Škorničková
jana_skornickova@nparks.gov.sg
1
Key Laboratory of Plant Resources Conservation
and Sustainable Utilization, Guangdong Provincial Key
Laboratory of Digital Botanical Garden, South China
Botanical Garden, Chinese Academy of Sciences, 723
Xingke Road, Tianhe District, Guangzhou 510650,
People’s Republic of China
2
University of Chinese Academy of Sciences, 19A,
Yuquan Road, Shijingshan District, Beijing 100049,
People’s Republic of China
3
Key Laboratory for Plant Diversity and Biogeography
of East Asia, Kunming Institute of Botany, Chinese Academy
of Sciences, Kunming 650201, People’s Republic of China
4
Horticulture Centre, South China Botanical Garden,
Chinese Academy of Sciences, Guangzhou 510650,
People’s Republic of China
5 oblong and smooth fruits, and pointed out that it was close
The Herbarium, Singapore Botanic Gardens, National
Parks Board, 1 Cluny Road, Singapore 259569,
Republic of Singapore
species in China and on original materials and methods, was
given in the first paper in this series and is not repeated
here (Ye et al. 2018). Since then, De Boer et al. (2018) have
recircumscribed Amomum as a monophyletic genus consist-
ing of species which have mainly winged or angled fruits
and fan-shaped or extended anther crests, and have recog-
nised additional five genera. According to this new treat-
ment, the Chinese species previously classified in Amomum
s.l. are now categorised into four genera namely Amomum
s.s. Roxb., Lanxangia M.F.Newman & Škorničk., Meistera
Giseke and Wurfbainia Giseke.
Hornstedtia tibetica T.L.Wu & S.J.Chen was originally
described by Wu and Chen (1978) based on a collection
from Medog County, Xizang (Tibet), China. The descrip-
tion, accompanied by a line drawing, was made from mate-
rial consisting of vegetative parts and infructescences only.
Wu and Chen (1978) placed this taxon in the genus Horn-
stedtia Retz. presumably due to the radical fusiform inflo-
rescences covered with rigid involucral bracts enclosing
to H. affinis Ridl. Without any subsequent collections of
flowering material, this taxon continued to be known only

13
Vol.:(0123456789)

1166
from fruiting material and remained in Hornstedtia in the
Flora Reipublicae Popularis Sinicae (Wu and Chen 1981)
and Flora of China (Wu and Larsen 2000).
During an expedition to the type locality of Hornsted-
tia tibetica in Medog County, Xizang in 2011, the third
author collected living material of this species and intro-
duced it to the South China Botanical Garden, Guangzhou,
where it has flowered in recent years. A close examination
of flowering material led to the suspicion that this species
was more closely related to Amomum than Hornstedtia. The
centre of the diversity of Hornstedtia is in evergreen South-
east Asia, with only a very few species extending through
monsoonal Southeast Asia to Southern China (Hainan and
Guangxi Province). The only Hornstedtia species known to
extend beyond this limit is H. arunachalensis S.Tripathi &
V.Prakash from near Dafla (Daphla) Hills (27°14′43.2″N,
93°42′20.0″E), which is only ca 300 km from the type local-
ity of H. tibetica in Medog County, China. Wu and Larsen
(2000) pointed out that H. arunachalensis might be conspe-
cific with H. tibetica.
The aim of this manuscript is to address the correct
generic placement of the species originally described as H.
tibetica and examine its relationship with H. arunachalensis.
In addition, we also examine the other Hornstedtia spe-
cies in China, H. hainanensis T.L.Wu & S.J.Chen, which
was also described from fruiting material only (Wu and
Chen 1978). Our recent collections, originating from the
vicinity of the type locality in Hainan Province, are success-
fully cultivated at South China Botanical Garden, Guang-
zhou. Study of the flowering material enables us to confirm
its correct generic placement in Hornstedtia as well as to
provide a detailed description and colour plate.
Materials and methods
Herbarium material
Type specimens of Hornstedtia tibetica, H. arunachalensis
and H. hainanensis from CDRI, IBK, IBSC, KUN and PE
(Thiers continuously updated) have been examined, data-
based and photographed. Additionally, six collections/nine
sheets of H. tibetica and 13 collections/15 sheets of H. hain-
anensis were examined from GXMI, IBK, IBSC, KUN and
PE.
Living material
Living material of H. tibetica and H. hainanensis was recol-
lected near their type localities and brought to cultivation
at South China Botanical Garden, where they flowered in
May 2014 and April 2015 and were vouchered as X.E.Ye
and L.Bai 14050701 (IBSC) and X.E.Ye and L.Bai 15040301
13
X.-E. Ye et al.
(IBSC), respectively. The vegetative parts as well as inflores-
cences and flowers were examined, dissected and measured.
The general plant terminology followed Beentje (2016),
while the description style followed recent works (e.g. Ye
et al. 2018, Lamxay and Newman 2012, Leong-Škorničková
et al. 2016). Pollen samples were collected from fresh flow-
ers and washed in 70% alcohol and further processed using
an ultrasonic cleaner (Branson 1510E-MT) for 18 min and
then centrifuged at 5000 r/s for 3 min. The supernatant was
removed, and the residue was re-suspended in 1 ml of 70%
alcohol. Washing followed by re-suspension was repeated
twice. Drops of pollen suspension were placed on sample
stubs, and then these were placed in the dryer for at least
1 day. The pollen grains are gold-coated in an auto-fine
sputter coater (JEOL JFC-1600) and observed under a high-
vacuum scanning electron microscope (SEM). Micrographs
were taken at 15 kV using the SEM. We measured the diam-
eter of 20 pollen grains, and then the average, maximum,
minimum and (P/E) ratio values were recorded to represent
the range of variation. The pollen terminology used followed
Kaewsri and Paisooksantivatana (2007). The method of pro-
cessing the pollen followed Ye et al. (2018).
Taxon sampling
Fifty-nine accessions representing 56 species were sam-
pled in this study. The origin of plant material together with
vouchers and accession numbers is presented in Table 1.
Of these, 16 Amomum s.l. species, Hornstedtia tibetica, H.
hainanensis and one Elettariopsis Baker species originated
from our collections in China. In addition, 40 ITS sequence
accessions were downloaded from GenBank including four
Amomum s.l. species, Hornstedtia arunachalensis and 30
species in the subfamily Alpinioideae to represent nine gen-
era of Alpinieae based on Kress et al. (2002), as well as all
Amomum s.l. and Alpinia clades or groups based on Xia
et al. (2004) and Kress et al. (2005). The choice of out-
groups, Tamijia flagellaris S.Sakai & Nagam. and Sipho-
nochilus aethiopicus (Schweinf.) B.L.Burtt followed Kress
et al. (2002).
DNA extraction
Total genomic DNAs of all species were extracted from
silica-gel-dried leaves using the CTAB method with minor
modification (Doyle and Doyle 1987).
DNA amplification and sequencing
The entire ITS1-5.8S-ITS2 region was amplified via poly-
merase chain reaction (PCR) using ITS5 and ITS4 (White
et al. 1990) with taq polymerase. The double-stranded PCR
products were amplified as described by Xia et al. (2004).
Taxonomic studies of Amomum (Zingiberaceae) in China II 1167

Table 1  Vouchers for sequence data for the analyses of ITS of Hornstedtia and related taxa
No. Taxon name Groups of Alpinia Geographic Location Voucher Coll. No. GenBank accession References
[after Kress et al. (Herb.) No.
(2005)]

Aframomum
1 A. angustifolium Madagascar Kress 92-3403 (US) AF478704 Kress et al. (2002)
(Sonn.) K.Schum.
2 A. daniellii (Hook.f.) Gabon Kress 99-6375 (US) AF478705 Kress et al. (2002)
K.Schum.
Alpinia
3 A. aenea B.L.Burtt & Alpinia III Sulawesi Argent 0016 (E) AY742351 Kress and Specht
R.M.Sm. (2006)
4 A. argentea Alpinia IVd Borneo L-95.0364 (CS) AY742337 Kress et al. (2005)
(B.L.Burtt &
R.M.Sm) R.M.Sm.
5 A. abundiflora Alpinia I Sri Lanka Weerasooriya (PDA) AY742334 Kress et al. (2005)
(Thwaites) Burtt &
R.M.Sm.
6 A. calcarata Roscoe Alpinia IVc China Kress 94-3657 (US) AF478710 Kress et al. (2002)
7 A. conchigera Griff. Alpinia II China Kress 00-6706 (US) AF478712 Kress et al. (2002)
8 A. elegans (C.Presl.) Alpinia V Philippines Kress 99-6412 (US) AF478713 Kress et al. (2002)
K.Schum.
9 A. foxworthyi Ridl. Alpinia IVe Philippines Kress 98-6293 (US) AF478714 Kress et al. (2002)
10 A. galanga (L.) Alpinia II Indonesia Kress 02-7213 (US) AF478715 Kress et al. (2002)
Willd.
11 A. intermedia Gag- Alpinia IVb Japan Kress 97-5780 (US) AF478716 Kress et al. (2002)
nep.
12 A. zerumbet (Pers.) Alpinia IVa China Liao 020704 (SCIB) AY742389 Kress et al. (2005)
B.L.Burtt &
R.M.Sm.
Amomum
13 A. compactum Sol. ex China, Yunnan, X.E.Ye and Y.H.Tong MF802556a
Maton cultivated TYH-204 (IBSC)
14 A. coriandriodo- China, Yunnan X.E.Ye 15053103 MF802560a
rum S.Q. Tong & (IBSC)
Y.M.Xia
15 A. glabrum S.Q.Tong China, Yunnan X.E.Ye 15051207E MF802563a
(IBSC)
16 A. koenigii J.F.Gmel China, Yunnan X.E.Ye 15051809 MF802558a
(IBSC)
17 A. longipetiolatum China, Hainan X.E.Ye 14051533G MF802564a
Merr. (IBSC)
18 A. menglaense China, Yunnan Xia-726 (HITBC) AY351996 Xia et al. (2004)
S.Q.Tong
19 A. maximum Roxb. China Xia-725 (HITBC) AY351995 Xia et al. (2004)
20 A. paratsaoko China, Guangxi X.E.Ye 14060501 MF802561a
S.Q.Tong & and (IBSC)
Y.M. Xia
21 A. petaloideum China, Yunnan Kress 95-5508 (US) AY769826 Kress et al. (2002)
(S.Q.Tong) T.L.Wu
22 A. purpureoru- China, Yunnan X.E.Ye and Z.L. Lin MF802566a
brum S.Q.Tong & 14040301 (IBSC)
Y.M.Xia
23 A. putrescens D.Fang China, Guangxi X.E.Ye 14052016 MF802568a
(IBSC)
24 A. quadratiolaminare China, Yunnan X.E.Ye and Z.L. Lin MF802557a
S.Q.Tong 14033102 (IBSC)

13

1168 X.-E. Ye et al.

Table 1  (continued)
No. Taxon name Groups of Alpinia Geographic Location Voucher Coll. No. GenBank accession References
[after Kress et al. (Herb.) No.
(2005)]

25 A. queenslandicum Australia Kmn 1428 (HITBC) AY352004 Xia et al. (2004)

R.M. Sm.
26 A. sericeum Roxb. China, Yunnan X.E.Ye 15051817 MF802565a
(IBSC)
27 A. subcapitatum Y.M. China, Yunnan X.E.Ye 15060601 MF802567a
Xia (IBSC)
28 A. subulatum Roxb. India, Mizoram MZU/BT/010 KX065426
29 A. tibeticum T.L.Wu Yarlung Zangbo- 92169 KJ872195
& S.J.Chen Brahmaputra
30 A. tibeticum T.L.Wu India, Nagaland 103604 KJ872194
& S.J.Chen
31 A. tibeticum T.L.Wu Yarlung Zangbo- 92156 KJ872193
& S.J.Chen Brahmaputra
32 A. tibeticum T.L.Wu China, cultivated X.E.Ye 14050701 MF802569a
& S.J.hen (IBSC)
33 A. tsaoko Crevost & China, Guangxi X.E.Ye 14060502 MF802562a
Lemarié (IBSC)
34 A. uliginosum Thailand 99-0474 (LYON) AY352008 Xia et al. (2004)
J.Koenig
35 A. villosum Lour. China, Guangxi X.E.Ye 14052005E MF802555a
(IBSC)
36 A. yunnanense China, Yunnan X.E.Ye 15060201 MF802559a
S.Q.Tong (IBSC)
Camptandra
37 C. parvula (King ex Thailand Kress 99-6328 (US) AF478730 Kress et al. (2002)
Baker) Ridl.
Elettariopsis
38 E. kerbyi R.M.Sm. Borneo Kress 96-5746 (US) AF478746 Kress et al. (2002)
39 E. monophylla (Gag- China, Hainan X.E.Ye 14051701 MF802570a
nep.) Loes. (IBSC)
40 E. smithiae Y.K.Kam Thailand Kress 99-6313 (US) AY352013 Kress et al. (2002)
41 E. stenosiphon Sarawak Kress 01-6847 (US) AF478748 Kress et al. (2002)
(K.Schum.)
B.L.Burtt &
R.M.Sm.
Etlingera
42 E. elatior (Jack) Malaysia Kress 94-4251 (US) AF478749 Kress et al. (2002)
R.M.Sm.
43 E. yunnanen- China Kress and Wood AY769846 Harris et al. (2008)
sis (T.L.Wu & 95-5511 (US)
S.J.Chen) R.M.Sm.
Hornstedtia
44 H. conica Ridl. Malaysia, Perak Ped. and Joh. 1082 AF414481 Pedersen (2004)
(C)
45 H. gracilis R.M.Sm. Cultivated Mood 996 AF41448 Pedersen 2004
46 H. hainanensis China, cultivated X.E.Ye 14062401 MF802571a
T.L.Wu & S.J.Chen (IBSC)
47 H. hainanensis China Kress 97-5769 (US) AF478766 Kress et al. (2002)
T.L.Wu a&
S.J.Chen
48 H. havilandii Cultivated W91p334 AF414479 Pedersen (2004)
(K.Schum.)
K.Schum.

13
Taxonomic studies of Amomum (Zingiberaceae) in China II 1169

Table 1  (continued)
No. Taxon name Groups of Alpinia Geographic Location Voucher Coll. No. GenBank accession References
[after Kress et al. (Herb.) No.
(2005)]

49 H. leonurus Sarawak Sakai 365 (KYO) AB097237 Sakai et al. (2001)

(J.Koenig) Retz.
50 H. minor (Blume) Sarawak Sakai 381 (KYO) AB097238 Sakai et al. (2001)
Valeton
51 H. reticulata Sarawak Sakai 387 (KYO) AB097236 Sakai et al. (2001)
(K.Schum.)
K.Schum.
52 H. sanhan Vietnam Newman 202 (E) AY769844 Harris et al. (2008)
M.F.Newman
53 H. scottiana Australia Johansen et al. C230 AF414480 Pedersen (2004)
(F.Muell.) (C)
K.Schum.
Renealmia
54 R. alpinia (Rottb.) Tropical America Kress 99-6407 (US) AF478778 Kress et al. (2002)
Maas
55 R. battenbergiana Tropical Africa Kress 94-5277 (US) AF478779 Kress et al. (2002)
Cummins ex Baker
Siliquamomum
56 S. tonkinense Baill. China Kress 00-6802 (US) AF478791 Kress et al. (2002)
Siphonochilus
57 S. aethiopicus (Sch- East Africa GH 00-134 (US) AF478792 Kress et al. (2002)
weinf.) B.L.Burtt
Tamijia
58 T. flagellaris S.Sakai Sarawak Kazuyuki S55 (KYO) AF478797 Kress et al. (2002)
& Nagam.
Vanoverberghia
59 V. sepulchrei Merr. Philippines Kress 95-5562 (US) AF478798 Kress et al. (2002)

ITS internal transcribed spacer

a
 Sequences newly published in this study. The other data were recently downloaded from GenBank

Data analysis in 50% majority rule consensus trees. Posterior probabilities

Sequences were edited and assembled by SeqMan II (Swindell
and Plasterer 1997). Multiple alignment of ITS regions was
performed manually with subsequent adjustments by BioEdit
version 7.2.5.0 (Hall 1999) and analysed in MrBayes version
3.2.6 (Ronquist and Huelsenbeck 2003). The consensus tree
was edited in FigTree version 1.4.3 (Rambaut 2009).
The Bayesian analyses which followed Middleton et al.
(2017) were run over 1,000,000 generations or longer until
the standard deviation of the split sequences reached val-
ues lower than 0.01. Thus, the ITS Bayesian inference took
2,000,000 generations to reach an average standard deviation
of 0.007548. The heat was set at 0.02, because the default
setting of 0.1 did not allow for satisfactory swaps between
chains; the substitution model used was a GTR with gamma-
distributed rate variation across sites, allowing for a proportion
of invariable sites and using relative burn-in discarding the
first 25% of sampled trees. The output trees were summarised
marked in the trees of between 0.98 and 1 are considered
strong support, of between 0.8 and 0.98 medium, and values
lower than 0.8 are considered poor (Droop 2012).
Results and discussion
Generic placement of Hornstedtia tibetica
Morphology: Hornstedtia, a genus of about 32 currently
accepted species and five accepted varieties (WCSP 2017),
has been distinguished from other basally flowering Alpin-
ioideae genera by the following characters (Valeton 1921;
Holttum 1950; Pedersen 2004): 1. The inflorescence is a
spike or a spike-shaped head, fusiform or cyathiform in
shape. 2. The outer involucral bracts are of a rather rigid
texture converging at the top and forming a narrow canal
from which one to a few flowers can protrude (fusiform type

13

1170 X.-E. Ye et al.

of inflorescence) or are stiffly erect with reflexed tips and the
interior bracts become shorter and softer gradually towards
the centre (cyathiform type of inflorescence). 3. The corolla
is shaped rather like a snake’s head or duck’s beak and usu-
ally has a broadly linear slipper-shaped labellum sometimes
with semi-lunate sidelobes or auricles covering the base of
the stamen, although species with spathulate labellum with
bilobed or entire apex but without auricles have also been
recorded (e.g. H. conica Ridl., H. gracilis R.M.Sm., H. hain-
anensis, H. sanhan M.F.Newman, H. bella Škorničk.). 4.
The labellum is mostly red or sometimes pink with white,
cream, pink or rarely yellow margin. The labellum may
rarely be yellow in the genus (e.g. H. tomentosa [Blume]
Bakh.f., H. pininga [Blume] Valeton), and occasional indi-
viduals with a white labellum in at least two pink-flowered
species have also been recorded (H. scottiana [F.Muell.]
K.Schum., H. sanhan) (Newman 1995; Larsen et al. 1999;
Lamb et al. 2013). 5. The anther ends either with a flat
rounded crest or a small rounded mostly recurved lobe above
the thecae or is crestless. 6. The fruit is always smooth, ellip-
soidal, obovoid or lenticular shape.

Fig. 1  Amomum tibeticum: a
habit, b young ligule, c grown
ligule, d single inflorescence
(scale 2 cm), e, f flower (top
view), g flower and dissection
(from left to right, from top
to down): bract, single flower
with bracteole, bracteole, calyx,
corolla lobes, labellum, stamen
with style, stigma and floral
tube in front and side views,
ovary with epigynous glands
(scale 1 cm). Photographs by
Y.S. Ye and L. Bai; based on
X.E. Ye and L. Bai 14050701

13
Taxonomic studies of Amomum (Zingiberaceae) in China II
Hornstedtia tibetica forms large clumps of usually stilt-
rooted leafy shoots to 1.2–2.5 m tall. The spindle-shaped
inflorescences are composed of brownish purple to maroon,
rigid and coriaceous, densely brown hirsute bracts. The
fertile bract supports a single flower, which emerges from
the tip of inflorescence and develops into a smooth ellipsoi-
dal fruit. All these characters match well with the current
definition of Hornstedtia. On the other hand, the morphol-
ogy of the inflorescences and fruits is quite diverse in the
polyphyletic Alpinioideae genera, and these characters of H.
tibetica fit well within the variability of some other genera
(e.g. Amomum s.l., Etlingera Giseke) as well. However, it is
the flower morphology which raises the principal question
about generic placement of this taxon. The large, thin and
broadly sub-orbicular white labellum with a bright yellow
centre, the stamen composed of a short and narrow filament,
and an anther with connective extending into a fan-shaped
anther crest (Fig. 1g) have never been observed in any Horn-
stedtia species, but are typical in Amomum s.s. (as delimited
by De Boer et al. 2018 and corresponding to A. maximum
clade, fide Xia et al. 2004). Its spindle-shaped spike is also
common in some Amomum s.l. species, although it is slightly
narrower than in most species so far known and the bracts
persist to fruiting stage (unlike the bracts of most Amomum
s.l. species which usually start rotting by the time the fruits
form). The smooth ellipsoidal fruit looks similar to that of
some Hornstedtia species, although it has to be noted that
the fruit shape in Amomum s.s. varies widely from winged
(most species; at least in the upper part) to ridged (e.g. A.
menglaense S.Q.Tong), grooved (e.g. A. sericeum Roxb.) or
angled (species previously classified as Elettariopsis).
In addition, pollen morphology has been previously
reported to be useful for classification and systematics
within Zingiberaceae due to the variation in size, shape
and ornamentation of the exine (Theilade et  al. 1993;
Fig. 2  Scanning electron micrographs of pollen Amomum tibeticum based on X.E. Ye and L. Bai 14050701: a pollen grain (scale 10 μm), b detail
of exine ornamentation (scale 5 μm). Photographs by L. Bai
1171
Saensouk et  al. 2009; Nurainas et  al. 2011). Nurainas
et al. (2011) observed 14 taxa of Hornstedtia using light
microscopy (LM) and scanning electron microscopy
(SEM) following the first record of the pollen morphol-
ogy of Hornstedtia species (i.e. H. hainanensis) by Liang
(1988). The results of both showed that Hornstedtia is
characterised by monad or monolete pollen with prolate,
spheroidal and ovoid shape and rugulate ornamentation
of the exine. Nurainas et al. (2011) indicated that pollen
morphology had been particularly significant in differenti-
ating Hornstedtia from Amomum s.l. (Liang 1988; Kaewsri
and Paisooksantivatana 2007), Cornukaempferia Mood &
K.Larsen (Saensouk et al. 2009) and Globba L. (Susanti
2009), all of which have the exine ornamented with spines
(echinate sculpturing), and Zingiber Mill. which has cer-
ebroid or reticulate ornamentation of the exine (Theilade
et al. 1993; Nurainas et al. 2011). Our investigations of the
pollen morphology of H. tibetica based on our collection
X.E. Ye and L. Bai 14050701 and H. hainanensis based on
X.E. Ye 14052301 reveal that the pollen of H. tibetica is
spheroidal or subspherical and has the exine ornamented
with spines (Fig. 2), while the pollen of H. hainanensis is
spheroidal or subspherical and has rugulate ornamentation
of the exine (Fig. 3). Spiny ornamentation of the exine
is common in Amomum s.s. and is present in all Chinese
species, where H. tibetica belongs according to its floral
morphology (as discussed above) as well as the molecular
evidence provided below.
Molecular data: After multiple sequence alignment, the
lengths of the nucleotide sequences were 597 bp for ITS
including 79 and 157 variable parsimoniously non-inform-
ative and informative sites, respectively. All species of
subfamily Alpinioideae clustered together and presented a
monophyletic lineage with strong support (posterior proba-
bility, PP = 1) confirming the findings of Kress et al. (2002).
13

1172
Fig. 3  Scanning electron micrographs of pollen Hornstedtia hainanensis based on X.E. Ye 14052301: a pollen grain (scale 20 μm), b detail of
exine ornamentation (scale 5 μm). Photographs by X.E. Ye
Analyses of the ITS region have been widely employed in
previous phylogenetic studies of Alpinieae (Xia et al. 2004;
Kress et al. 2005, 2007; Droop 2012). We have therefore
employed this region in the present study to gain insight into
the phylogenetic placement of H. tibetica and H. arunacha-
lensis based on our samples enriched with available ITS
sequences from GenBank.
Xia et al. (2004) analysed 23 species of Amomum s.l.
using ITS and matK DNA sequences and resolved three
separate groups (namely the Amomum maximum group, the
A. villosum group and the A. tsaoko group).
Kress et al. (2007) performed combined ITS and trnK/
matK phylogenetic analyses of tribe Alpinieae. They
retrieved the same three distinct groups proposed by Xia
et al. (2004) and furthermore recognised seven well-sup-
ported lineages which corresponded to formerly recognised
genera (namely Aframomum K. Schum., Renealmia L. f.,
Alpinia galanga clade, Etlingera, Geocharis [K. Schum.]
Ridl., Geostachys [Baker] Ridl. and Siliquamomum Baill.).
They also retrieved another seven well-supported lineages
(Alpinia fax clade, Alpinia eubractea clade, Amomum vil-
losum clade, Alpinia carolinensis clade, Alpinia zerumbet
clade, Amomum tsaoko clade and Alpinia rafflesiana clade)
and three problematic lineages (Amomum maximum clade,
Elettariopsis clade and Hornstedtia grade). Within the
problematic lineages, they pointed out that seven species
of Hornstedtia sampled formed a paraphyletic group nest-
ing with Etlingera and that the Elettariopsis clade which
contained species of Elettariopsis, Amomum s.l. and Para-
momum S.Q.Tong was only moderately supported.
Droop (2012) focused in her PhD work on the phylogeny
of Amomum s.l. and conducted phylogenetic analyses of ITS
sequence data, which were more extensive in sampling than
those of Xia et al. (2004) and Kress et al. (2007). Other than
the three Amomum clades of Xia et al. (2004), five additional
13
X.-E. Ye et al.
groups of Amomum s.l. were also observed in her ITS tree.
The genera Aframomum, Alpinia, Elettaria Maton, Elettario-
psis, Etlingera, Leptosolena C.Presl, Geocharis, Geostachys,
Plagiostachys Ridl., Renealmia, Vanoverberghia Merr., and
species in Hornstedtia are nested in these groups. These
results are in agreement with Xia et al. (2004) and Kress
et al. (2007).
De Boer et al. (2018) have further investigated the phy-
logeny of Amomum s.l. using matK and nrITS. Amomum s.l.
is shown to consist of nine clades and Alpinia of six. The
genera Elettaria, Elettariopsis, Etlingera, Geocharis, Geo-
stachys, Plagiostachys and species in Hornstedtia are nested
in these clades. Morphological studies of species previously
subsumed in Amomum s.l. supported the recognition of new
genera that corresponded to well-delimited clades in the
phylogenetic framework and led to the resurrection of three
genera, Conamomum Ridl., Meistera Giseke and Wurfbainia
Giseke, and the description of three new genera, Epiamo-
mum A.D.Poulsen & Škorničk., Lanxangia M.F.Newman &
Škorničk. and Sundamomum A.D.Poulsen & M.F.Newman.
Although the sampling in our study is less extensive than
that of Kress et al. (2007), Droop (2012) and De Boer et al.
(2018), the material selected includes representatives from
previously recognised major lineages in order to establish
the generic placement of H. tibetica. In general, our result
is congruent with all studies mentioned above in terms of
retrieving previously recognised lineages and their topology
(Fig. 4). We have recognised five well-supported lineages
which correspond to formerly recognised genera (namely
Aframomum [PP = 1], Renealmia [PP = 1], Alpinia galanga
clade [PP = 1], Etlingera [PP = 0.99] and Siliquamomum),
six well-supported lineagestaxonomic names (Alpinia fax
clade, Alpinia eubractea clade [PP = 1], Amomum villo-
sum clade/Wurfbainia [PP = 0.928], Alpinia carolinensis
clade, Alpinia zerumbet clade [PP = 1] and Amomum tsaoko
Taxonomic studies of Amomum (Zingiberaceae) in China II
Fig. 4  The 50% majority rule consensus tree resulting from the Bayesian inference of the ITS data. Statistical support is expressed as posterior
probabilities, which are shown by the nodes
clade/Lanxangia [PP = 1] and the two problematic line-
ages (Amomum maximum clade/Amomum s.s. [PP 0.999] and
Hornstedtia grade [PP = 0.971]). Our study also confirms
that several genera in Alpinioideae, including Amomum s.l.
and Hornstedtia, are polyphyletic. The type of Hornstedtia,
H. scyphifera (J.König) Steud., was also not included in the
above studies, but based on the morphology it is predicted
1173
to fall into the clade with H. reticulata, which is morpho-
logically very similar to H. scyphifera that was previously
included in the work of Droop (2012).
We have therefore included 11 morphologically diverse
Hornstedtia species including H. reticulata. Our results
concur with previous studies (Kress et  al. 2007, Droop
2012) that Hornstedtia is polyphyletic and, as found by
13

1174
Kress et al. (2007), all sub-clades containing Hornstedtia
species fall within another well-supported (PP = 0.9714)
clade with Etlingera, Alpinia eubractea clade and Vanover-
berghia, which is sister to the well-supported (PP = 0.928)
Amomum villosum clade/Wurfbainia (as per Xia et al. 2004,
Kress et al. 2007 and De Boer et al. 2018). It is clear that H.
tibetica is not affiliated to any of the sub-clades containing
Hornstedtia species.
In our analysis, Hornstedia tibetica and H. arunacha-
lensis are both nested in the Amomum maximum clade
which has been recircumscribed as Amomum s.s.(De Boer
et al. 2018), and these two species form a well-supported
sub-clade with strong support (PP = 1).
Based on the morphological and molecular evidence
discussed above, we conclude that Hornstedia tibetica
and H. arunachalensis belong to Amomum s.s.
Relationship between Hornstedtia tibetica and H.
arunachalensis
It remains unclear why Tripathi and Prakash (1999)
decided to place Hornstedtia arunachalensis in Hornst-
edtia and compared it to H. scyphifera (J.Koenig) Steud.
because the floral features of H. arunachalensis (namely
the large white labellum and the anther with prominent
fan-shaped crest) are not known to occur in Hornstedtia.
Using the original description and the illustration of
Hornstedtia arunachalensis, Wu and Larsen (2000) were
the first to suggest that it might be conspecific with H.
tibetica, but, without flowering material of H. tibetica,
they were unable to make a formal decision.
As mentioned above, Hornstedtia tibetica has large
white flowers and, after detailed comparison of multiple
morphological characters (Table 2), we believe that they
represent the same taxon. Even though there are slight
differences in the density of indumentum on the peti-
ole and floral tubes and the sizes of various floral parts
seem to be slightly smaller in H. arunachalensis, we do
not regard these differences as sufficiently significant to
warrant maintaining H. tibetica and H. arunachalensis as
two distinct taxa (Table 2). Particularly, the differences in
sizes are quite likely caused by comparing fresh and dry
material, while the indumentum density is often influ-
enced by age (i.e. certain parts become glabrous with
age).
The type localities of Hornstedtia arunachalensis and
H. tibetica are about 300 km apart. The geographic prox-
imity and pattern of all three populations of H. arunacha-
lensis and five of H. tibetica, as well as the similar alti-
tude and ecological preferences, also suggest that they
represent a single species.
13
X.-E. Ye et al.
Furthermore, our ITS analyses indicate (as shown in
Fig. 4) that Hornstedtia tibetica and H. arunachalensis
cluster together in the Amomum maximum clade forming a
single strongly supported group (PP = 1), without forming
two clearly separate sub-clades, further supporting our
decision to treat them as conspecific.
As a result, in the following taxonomic treatment, we
transfer Hornstedtia tibetica into Amomum s.s. and treat
H. arunachalensis as a synonym of it.
Taxonomic treatment
Amomum tibeticum (T.L.Wu & S.J.Chen) X.E.Ye, L.Bai
& N.H.Xia, comb. nov. (Figs. 1, 2).  ≡ Hornstedtia tibetica
T.L.Wu & S.J.Chen, Acta Phytotax. Sin. 16 (3): 39. 1978;
T.L.Wu & S.J.Chen, Wu (ed.) Fl. Reipubl. Popularis Sin.
16(2): 136. 1981; T.L.Wu & K.Larsen, in Wu and Raven
(eds.) Fl. China. 24: 58. 2000.—LECTOTYPE (designated
here): China. Xizang Province: Medog County, Beibeng
Township (previously District), 810 m a. s. l., 11 Aug 1974,
Qingzang expedition 74-1913 (PE! barcode PE00075268);
isolectotypes: PE PE00075267!, KUN KUN1219292!,
KUN1219293!).
=  Hornstedtia arunachalensis S.Tripathi & V.Prakash,
Nordic J. Bot. 19: 329. 1999. syn. nov.—TYPE: near Dafla
(Daphla) Hills (27°14′43.2″N, 93°42′20.0″E) (cited in
the protologue as “Arunachal Pradesh State, Papum Pare
[Papampara] Dt., Umpham”), 1 May 1997, Sunil Tripathi
20849 (holotype: CDRI No.32200!; isotype: CAL [n.v.]).
Herb 1.2–2.5 m tall, clump-forming, with 7–12 pseudostems
per clump; rhizomes scarlet externally, stilt roots present, scar-
let. Leafy shoot composed of 11–23 leaves per pseudostem;
pseudostem slightly swollen at base, ca 3 cm diameter, scar-
let at base, and green with brownish red tinge distally; leaf
sheaths green with red tinge on the margin, densely hirsute
when young, turning glabrous with age; ligule narrowly
elliptic to ovate, usually entire, sometimes emarginate in the
upper leaves, ca 1.3 cm long, red or green with red tinge and
densely ciliate on the margin when young, turning brownish
black, thin coriaceous to coriaceous, turning dry and glabrous
with age; petiole 0.5–1 cm long, densely hirsute when young,
turning glabrous with age, green; lamina linear to narrowly
oblong, 54–60 × 6.5–11 cm, green and glabrous above, yel-
lowish green and densely brownish hirsute below, base cune-
ate, apex caudate, membranous, margin entire. Inflorescence
radical, entirely above ground; peduncle up to 1 cm long,
8 mm diameter, covered by sheathing scales; scales ca 6 in
number, ovate or broadly ovate at base (ca 1 × 1 cm) to nar-
rowly ovate distally (ca 14 × 6 mm) when flattened, brown-
ish purple to maroon, pink at base, coriaceous, glabrous
adaxially, densely brown hirsute abaxially, apex emarginate,
rigid, mucronate abaxially; flowering part narrowly fusiform,
 Table 2  Comparison of the main morphological characters between Amomum tibeticum (based on type, protologue and living flowering material of X.E. Ye and L. Bai 15040301) and Hornsted-
tia arunachalensis (based on type and protologue)
Characters Amomum tibeticum Hornstedtia arunachalensis

Herb 1.2–2.5 m tall Up to 2 m tall

Ligule Narrowly elliptic to ovate, ca 1.3 cm long, densely ciliate on the margin when young, Narrowly elliptic, ca 1.5 cm long, subcoriaceous, glabrous
turning glabrous with age
Taxonomic studies of Amomum (Zingiberaceae) in China II

Petiole 0.5–1 cm long, densely hirsute when young, turning glabrous with age
Lamina Linear to narrowly oblong, 54–60 × 6.5–11 cm, green, glabrous above, densely brown-
ish hirsute below, base cuneate, apex caudate, margin entire
Spike Narrowly fusiform, 9.5–16.5 × 2.4–3 cm; 1 or 2 flowers open at once
Calyx 5–6 cm long, white with slight pink tinge, pubescent, apex bilobed or trilobed
Floral tube ca 8.5 cm long, creamy white, densely or sparsely pubescent
Corolla lobes Dorsal oblong, ca 4 × 1.7 cm, white, apex rounded and cucullate, lateral narrowly
oblong to narrowly ovate, ca 4 × 1.2 cm, white, glabrous
Labellum Sub-orbicular, 4–5 × 3.5–4 cm, white with central bright yellow band, pubescent on
the band above; margin revolute and crisped, apex rounded
Stamen Anther thecae ca 1.7 cm long, with densely glandular hairs; anther crest fan-shaped
to obscurely trilobed, slightly revolute, truncate broadest at apex, ca 6 mm long, ca
13 mm wide, white
Ovary Cylindrical, ca 14 × 5 mm, brown pubescent at base, pinkish yellow pubescent on the
upper part
Style Filiform, ca 10 cm long, glabrous, stigma cup-shaped, dorso-ventrally pressed, gla-
brous, ostiole ciliate
1 cm long, pubescent
Narrowly elliptic, up to 60 × 8 cm, dark green, glabrous above, puberulous below, midrib
with long hairs, base cuneate, apex caudate, margin entire
Fusiform with narrower, acute tip, ca 10 × 3 cm; 2 flowers open at once
ca 5 cm long, white, sparsely pubescent, bilobed (trilobed sometimes)
ca 8 cm long, pink, glabrous
Dorsal oblong ca 3.5 × 1.4 cm, whitish, apex rounded; ca 3.5 cm long; lateral narrowly
oblong to narrowly ovate, ca 3.5 × 0.7 cm, white, glabrous
Obovate, ca 4 × 2.5 cm, whitish, slightly pubescent above, margin crenulate, apex
rounded
Anther thecae ca 1.5 cm long, slightly hairy; anther crest semi-lunate to trilobed, trun-
cate broadest at apex, ca 4 mm long, ca 10 mm wide, yellowish
Cylindrical, ca 10 × 3 mm, villous
Filiform, ca 10 cm long, glabrous, stigma cup-shaped, dorso-ventrally pressed, ostiole
ciliate

Differences in bold

13
1175

1176
9.5–16.5 × 2.4–3 cm (excluding flowers), brownish purple to
maroon; 1 or 2 flowers open at once; bracts 14–30 in num-
ber, narrowly ovate to narrowly oblong, 10–11.5 × 1.8–2.6 cm,
white with pink tinge at apex, thin coriaceous, subtending a
single flower, apex acute, margin ciliate towards apex; bracte-
ole linear, ca 5.5–7.6 × 0.7–1 cm, keeled, semi-translucent to
creamy white, appressed pubescent, apex acuminate. Flower
ca 15 cm long including pubescent, ca 4-mm-long pedicel;
calyx 5–6 cm long, white with slight pink tinge, pubescent
externally, glabrous internally, apex bilobed or trilobed; lobes
ca 1.5 × 2.7 cm long; floral tube cylindrical, slightly widened
at the throat, ca 8.5 cm long, ca 3 mm diameter, creamy white,
densely or sparsely pubescent externally and internally; dor-
sal corolla lobe oblong, ca 4 × 1.7 cm, white, apex rounded
cucullate, glabrous; lateral corolla lobes narrowly oblong to
narrowly ovate, ca 4 × 1.3 cm, white, glabrous, apex cucullate;
labellum sub-orbicular, 4–5 × 3.6–4 cm, white with bright yel-
low central band, band 3.5–4 cm × 3.6–4 mm, pubescent on
the band above, glabrous below and with transparent vena-
tion radiating towards margin; margin prominently revolute
and crisped; lateral staminodes vestigial, white with red or
pink tinge or absent; stamen white, ca 2.5 cm long; filament
flat, ca 2 mm long, 3 mm wide, with glandular hairs; anther
oblong, 2.3 cm long (including straightened crest), ca 4 mm
wide, with dense glandular hairs; anther crest fan-shaped,
obscurely trilobed, slightly revolute, ca 6 mm long, ca 13 mm
wide, membranous, white, glabrous; anther thecae two, par-
allel, dehiscing along entire length; pollen grains spherical
or subspherical, 57.64 (45.98–65.52) μm, exine sculpture
(ornamentation) echinate, uniformly distributed, 2–3  µm
tall, 2–4 µm thick at base; epigynous glands two, subulate,
14 mm long, 2 mm diameter at base, apex acuminate, pale
yellow, glabrous; ovary cylindrical, ca 14 × 5 mm, brown
pubescent at base, pinkish yellow pubescent on the upper part,
trilocular, placentation axile; style filiform, ca 10 cm long,
white, glabrous; stigma cup-shaped, dorsiventrally pressed,
glabrous, ostiole ciliate, facing upwards. Infructescence
peduncle 8–15 mm. Fruit ellipsoidal, 2.6–3 × 1.2–1.5 cm,
smooth (without any kind of protuberance), pale pink, pale
yellow pubescent, apex with persistent 3–5-cm-long calyx;
seeds ellipsoidal, ca 2.1 × 1.7 mm, brownish, glabrous; aril
membranous, semi-translucent.
Chinese name: This species was originally proposed as 西
藏大豆蔻 (romanised xi zang da dou kou) in the protologue.
The Chinese name 大豆蔻 (da dou kou) generally refers to
Hornstedtia species, but, with the change in generic place-
ment, we propose to change the Chinese botanical name into
西藏豆蔻 (xi zang dou kou) .
Phenology: Flowering starts from late March, and fruiting
has been recorded in August.
13
X.-E. Ye et al.
Distribution area: So far known to occur from China (S
Xizang) to India (Nagaland).
Habitats and ecology: According to our observations, infor-
mation obtained from herbarium specimens and the litera-
ture (Wu and Chen 1978, Tripathi and Prakash 1999), this
species occurs at an altitudinal range of 650–1300 a.s.l., usu-
ally on hillsides in the understorey of broad-leaved forest,
sometimes on shady stream banks.
Provisional IUCN conservation status: This species has a
large extent of occurrence (around 21,556 km2), and it is
therefore here proposed as globally of Least Concern (LC)
according to the latest IUCN Standards and Petitions Sub-
committee (2017) guidelines. Not all of this area is, however,
suitable for this species, and the habitat in certain parts is
degrading. None of the eight currently known localities is
in a protected area. It is therefore possible that with a better
understanding of AOO and population sizes, A. tibeticum
may need to be reassessed as vulnerable.
Vernacular names and uses: According to the collecting
information on Qingzang expedition 74-4483 (PE, KUN),
the plant is known as “Su-Men” by the local Monba ethnic
minority (Xizang, China), who eat its seeds. The plant is
also known as “B-lak” where the flowers are used as orna-
mentation in marriage ceremonies by Adi tribal women (Tri-
pathi and Prakash 1999).
Notes: Wu and Chen (1978) designated the collection
Qingzang expedition 74-1913 deposited at PE as the type
of Hornstedtia tibetica. We have located two separate
sheets of this collection at PE (barcoded as PE00075268
and PE00075267), of which we select the sheet consisting
of two infructescences and a single leaf as the lectotype
(PE 00075268). The second sheet at PE (PE00075267),
as well as two additional sheets located at KUN (barcoded
as KUN1219292, KUN1219293), shall be regarded as
isolectotypes.
Additional specimens examined:  CHINA. Guangdong:
cultivated in South China Botanical Garden, Guangzhou, 7
May 2014, X.E. Ye and L. Bai 14050701 (IBSC); Xizang:
Nyingchi City: Medog County, 1000 m a. s. l., 23 Aug 1974,
Qingzang expedition 74-4483 (PE00075269, PE00075270,
KUN0335795, KUN0335796); Medog County, Beibeng
Township, Jiangxin Village, 850 m a. s. l., 3 Jun 1983, B.S.
Li and S.Z. Cheng 04988 (PE01314799); Medog County,
Beibeng Township, Bangu Mountain, 1300 m a. s. l., 16
Jul1980, Plateau Group, Department of Ecology 11132
(PE01314798); Medog County, Beibeng Township, 650 m
a.  s.  l., 14 Sep 1974, Anonymous 2546 (PE01790340,
PE01790341); Medog County, Y.S. Ye 5630 (IBSC).
Taxonomic studies of Amomum (Zingiberaceae) in China II 1177

Fig. 5  Hornstedtia hainanensis: a habit, b clustered inflorescences, c, labellum, ovary with epigynous glands, stamen in front view, and flo-
d flowers in front and side views, e single inflorescence (scale 1 cm), ral tube with stamen in back view (scale 1 cm). Photographs by Y.S.
f flower and dissection (from left to right, from top to down): bract, Ye and L. Bai; based on X.E. Ye and L. Bai 15040301
bracteole, calyx, single flower with bracteole removed, corolla lobes,

Hornstedtia hainanensis T.L.Wu & S.J.Chen, Acta Phy- (Figs.  3, 5).—LECTOTYPE (designated here): China,
totax. Sin. 16 (3): 38. 1978; T.L.Wu & S.J.Chen, Wu (ed.) Hainan Province, Sanya City (Ya Zhou), Yanglin, ca
Fl. Reipubl. Popularis Sin. 16(2): 136. 1981; T.L.Wu 450  m a.  s.  l., 6 Mar 1933, F.C.How 70318 (IBSC
&K.Larsen, Wu and Raven (eds.) Fl. China. 24: 58. 2000

13

1178
barcode IBSC0005277!; isolectotypes: IBSC0005278!,
IBK00137677!, PE00075318!).
Herb 1.1–2.3 m tall; rhizome near soil surface. Leafy shoot
composed of 9–20 leaves per pseudostem; pseudostem
slightly swollen at base, ca 2.5 cm diameter, brownish at
base, yellowish green at apex; leaf sheaths yellowish green,
glabrous; ligule entire, 7–9 mm long, thin coriaceous to
coriaceous, green with brown tinge, tomentellous at junc-
tion with sheath, margin ciliate, green when young, with
a narrow dark or pale brown stripe when mature; petiole
nearly none to ca 0.5 cm long; lamina linear to narrowly
oblong, 30–50 × 5–7  cm, green above, yellowish green
below, coriaceous, base cuneate or rounded, apex caudate,
glabrous on both sides. Inflorescence radical, entirely above
ground; peduncle to 5 mm long, covered by sheathing scales;
scales ca 12 in number, broadly ovate and 2-ranked at base
(ca 1 × 3.5 cm) to ovate distally (ca 6 × 5 cm), carmine api-
cally, pink basally, coriaceous, yellowish white appressed
pubescent externally, glabrous internally, margin ciliate,
apex emarginate, rarely rigid mucronate; flowering part
narrowly ovoid, 9–11 × 3–4 cm (including flowers), pubes-
cent at base; fertile bracts ca 24 in number, each subtending
a single flower, pale shiny pink, semi-translucent at base,
narrowly triangular, 5.7–6.5 × 1.7–3 cm, thin coriaceous,
glabrous on both sides; bracteole absent. Flower ca 8 cm
long, floral tube rarely much exserted beyond the bracts at
anthesis, but further elongates additional 2–3 cm after anthe-
sis; calyx tubular at base, 4.2–4.8 cm long, unilaterally split
to ca 1.2 cm, upper part (ca 2.7 cm) widened, pubescent
externally, glabrous internally, apex tridentate; floral tube
cylindrical, slightly widened at the throat, 5.2–5.5 cm long
when stretched, 2.5 mm diameter, white, pink tinged at apex,
pubescent externally and internally; dorsal corolla lobe
oblong, ca 16 × 8 mm, apex rounded and cucullate, pink, gla-
brous; lateral corolla lobes narrowly oblong, ca 13 × 5 mm,
apex rounded and cucullate, pink, glabrous; labellum spathu-
late, ca 2.2 × 1.2 cm, pubescent above, glabrous below except
slightly pubescent at base, basal part thick and fleshy, ca
6 mm wide, pale pink, tinged with two darker pink bands,
upper part ca 1.2 cm wide, membranous, pink, margin prom-
inently crisped and fimbriate, apex bifid; lateral staminodes
absent; anther sessile, ca 12 × 3 mm, with dense glandular
hairs at sides, long pubescent; pollen grains creamy white,
ellipsoidal, 90.6 (76–107.5) × 84.1 (72.5–100) μm, exine
sculpture (ornamentation) psilate under LM and rugulate
with long ridges under SEM, without aperture; anther crest
absent; anther thecae two, parallel, upper 1–2 mm dehis-
cent, lower part adnate; epigynous glands 3, forming a tube,
flat, two of them slightly longer, ca 3 mm long, shorter one
ca 2 mm long, creamy white, glabrous; ovary cylindrical,
8 × 3 mm, pubescent, trilocular, placentation axile; Style fili-
form, white, pubescent; stigma obtrigonous, ostiole broad,
13
X.-E. Ye et al.
transversely elliptic, ciliate, mucilaginous, facing ventrally.
Fruit ellipsoidal, 2–2.5 × 1.2–1.6  cm, pericarp smooth
(without any kind of protuberance), membranous, semi-
translucent, glabrous, apex with persistent 4–4.5-cm-long
calyx; seeds ellipsoidal; aril membranous, semi-translucent.
Chinese name: 海南大豆蔻 (romanised by hai nan da dou
kou) (this new name which indicates the locality is sug-
gested. The previous name 大豆蔻 [da dou kou] generally
refers to Hornstedtia species).
Phenology: Flowering in early March–April and fruiting in
March–August.
Distribution area: So far known only from China (Guangxi,
Guangdong and Hainan).
Habitats and ecology: This species has been reported to
grow usually on hillsides in the understorey of broad-leaved
forest, often near streams. It can be found at 500–600 m a.s.l.
Provisional IUCN conservation status: This species has
a large extent of occurrence (around 21,793 km2) and is
known from more than 10 localities some of which are pro-
tected. We, therefore, propose this species to be treated as of
Least Concern (LC) according to the latest IUCN Standards
and Petitions Subcommittee (2017) guidelines.
Vernacular names and uses: The species is called “烂头
溶 (romanised lan tou rong)”, “烂头羊 (lan tou yang)” or “
烂头包 (lan tou bao)” by the local Zhuang ethnic minority
(Guangxi, China), who eat its sour and sweet fruits accord-
ing to the information provided on the labels of D.Y.Liu
76638, X.P.Liao and D. Fang 76614, and D.Fang 76549 (all
deposited at GXMI). The whole plants are also used as a
medicinal plant for heat-clearing and detoxicating according
to information from D.Fang 76549 (GXMI).
Notes: In the protologue, the collection of F.C.How 70318
deposited at IBSC was designated as the type of Hornstedtia
hainanensis. We have located two separate sheets of F.C.How
70318 at IBSC (IBSC0005277, IBSC0005278). Of these, we
select the sheet with the base of the plant, two leaves, two
infructescences and three fruits (IBSC0005277) as the lecto-
type of H. hainanensis. The remaining sheet (IBSC0005278)
and another two sheets located at PE and IBK (PE00075318,
IBK001137452) are to be treated as isolectotypes.
Although H. hainanensis was originally described from
fruiting material only, current morphological and molecular
evidence as well as palynological evidence from the mate-
rial recollected near the type locality confirms the correct
generic placement of this taxon.
Additional specimens examined: CHINA. Guangdong:
Yangchun County, Baiyong Nature Reserve, 600  m
a. s. l., 21 May 1991, N. Liu 1842 (IBSC0022897); culti-
vated without detailed locality, 15 Mar 1982, T.L. Wu and
Taxonomic studies of Amomum (Zingiberaceae) in China II
S.J. Chen 4 (IBSC0022902); Guangxi: Fangchenggang
City, Shangsi County, Nadang Township, Sancha River,
29 May 1982, T.L. Wu and S.J. Chen 11 (IBSC0022901);
Fangchenggang City, Dongxing County, Tansan Town-
ship, Lihuo Village, Dazha, 27 Apr 1977, D.Y. Liu 76638
(GXMI 045299, GXMI 045301); ibid., 23 May 2014, X.E.
Ye 14052301 (IBSC); Dongxing County, Tansan Town-
ship, Lihuo Village, Caiwen Mountain, 500 m a. s. l., 18
Aug 1975, X.P. Liao and D. Fang 76614 (GXMI 045300);
Dongxing County, Nakan Township, Laowuchang, 8 May
1971, D. Fang 76549 (GXMI 045302); Hainan: Wuzhis-
han City, Nansheng Town (previously Ding’an County,
Tongjia), 1 Aug 1934, X.R. Liang 64268 (Paratype)
(IBSC0022896, IBK00137452); Wanning City, Xinglong
District, Shenjingling, 333 m a. s. l., 10 Apr 1935, F.C.
How 71789 (paratype, IBSC0022895); Baisha County,
Yuanmentong, 10 Apr 1936, X.Q. Liu 26088 (paratype,
IBSC0022894); Lingshui County, Sanjiao Ling, 16 Jan
1934, C. Wang 36766 (paratype, IBSC0022899); Sanya
City, Xiaoqiangkang, 19 Jul 1933, X.R. Liang 62202
(paratype, IBSC0022900); Dongfang County (previously
Gan’en County), 20 Feb 1934, X.R. Liang 64979 (para-
type, IBSC0022898).
Acknowledgements  We thank the curators of CDRI, GXMI, IBK,
IBSC, KUN, SING and PE herbaria for permission to examine and
photograph specimens in their care. The first author would like to thank
Peng Feng, Cheng Yu-Fen (IBSC) and Dr. Chen Juan (IBSC) for assis-
tance in imaging specimens from IBSC and PE; Hu Xiao-Ying (SCBG)
is acknowledged for assisting with the SEM pollen scanning. We thank
Dr. Carmen Puglisi (SING) for guidance on molecular analysis and Dr.
Nurainas (ANDA) for kindly providing her papers. X.-Y. Ye is grate-
ful to the University of Chinese Academy of Sciences and Singapore
Botanic Gardens for supporting the study at SING. The research of
X.-Y. Ye and N.-H. Xia was supported by the National Natural Sci-
ence Foundation of China (Grant no. 31170185); the research of the
last author is supported by National Parks Board, Singapore, and by
the Czech Science Foundation, GAČR (Grant No. P506/14/13541S).
We thank Dr. Mark Newman for extensive language revision of this
manuscript.
Compliance with ethical standards  Liang YH (1988) Pollen morphology of the family Zingiberaceae in
Conflict of interest  The authors declare that there is no financial or
non-financial conflict of interest.
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