全 文 ：植物生态学报 2008, 32 (4) 751~759
Journal of Plant Ecology (Chinese Version)

——————————————————
Received: 2006-09-20 Accepted: 2007-08-11
Foundation item: the National Natural Science Foundation of China (30170160)
Thanks for the field help of Ling Zhang, Fan Chen, Xing-Sheng Yu, Xiu-Xia Gao, Wen-Biao Li, Qiao-Ming Li and Yong Liu, for the writing help of
Quan-Guo Zhang and Matthew Warren, and for the lab assistance of the Molecular Phylogenetics and Biogeography Lab of Kunming Institute of Botany,
Chinese Academy of Sciences
*Author for correspondence E-mail: cj@xtbg.org.cn
具混合繁殖策略的草本植物异果舞花姜
的居群遗传结构
周会平1,3 陈进1* 张寿洲2
(1 中国科学院西双版纳热带植物园,云南勐腊 666303) (2 深圳仙湖植物园,广东深圳 518004)
(3 中国科学院研究生院,北京 100049)
摘 要 居群遗传结构的形成受到各种因素的影响。其中, 繁殖方式可能对居群内遗传变异有极其重要的意义,
而距离隔离也是居群间变异产生的主要原因之一。异果舞花姜(Globba racemosa)具有混合繁殖策略(以种子进行有
性繁殖和以珠芽进行无性克隆繁殖)。调查分布于云南的7个异果舞花姜居群间有性与无性克隆繁殖的差异。采用
ISSR标记研究各个居群的遗传多样性与克隆多样性, 探讨繁殖方式和距离隔离对居群遗传结构的影响。调查结果
表明, 异果舞花姜各个居群存在一定的繁殖差异。ISSR结果显示, 该种在种水平上呈现较高水平的遗传变异
(PPB=71.19%), 大部分的变异来自于居群间(GST = 0.590 7)。同时, 异果舞花姜具有较高水平克隆多样性(G/N =
0.88)。遗传多样性和克隆多样性与繁殖水平的变异间相关性不明显, 说明繁殖方式不是居群遗传结构形成的必要
和决定性的因素。居群间的地理距离与遗传距离显著相关(r = 0.68, p < 0.05), 表明距离隔离是居群间遗传变异形
成的重要原因。其它因素(如少量新有性个体的补充、细胞突变、奠基效应等)也对异果舞花姜居群遗传结构的形
成和维持起到了重要作用。
关键词 异果舞花姜 有性繁殖 克隆繁殖ISSR 遗传多样性 克隆多样性
GENETIC AND CLONAL DIVERSITY OF GLOBBA RACEMOSA, A HERB WITH
A MIXED REPRODUCTIVE MODE
ZHOU Hui-Ping1,3, CHEN Jin1*, and ZHANG Shou-Zhou2
1Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, Yunnan 666303, China, 2Shenzhen Fairylake Botanical
Garden, Shenzhen, Guangdong 518004, China, and 3Graduate University of Chinese Academy of Sciences, Beijing 100049, China
Abstract Aims Our objective was to understand the relative effects of reproductive mode, geo-
graphic distance and other potential factors on within- and among-population genetic and clonal diver-
sity in Globba racemosa, a species that can reproduce sexually (via seeds) and asexually (via bulbils).
Methods We investigated reproduction variation in the seven natural G. racemosa populations in
Yuannan Province, China by measuring the number of sexually produced fruits and asexually produced
bulbils. Genetic data were collected using inter-simple sequence repeat (ISSR) markers. We evaluated
the distribution of genetic variance and levels of population genetic diversity and assessed the relation-
ship between genetic diversity and reproduction variation.
Important findings Globba racemosa has a relatively high genetic diversity at the species level
(PPB=71.19%), and most genetic variances resided among populations (GST = 0.590 7). A high clonal
diversity was found in this species (G/N = 0.88). No significant difference was detected between repro-
ductive mode and genetic diversity or clonal diversity, which suggested that the mode of reproduction
was not necessarily a determinant of population genetic diversity. There was a significant correlation
between genetic and geographical distances (r = 0.68, p < 0.05), which indicated that genetic
differentiation between populations was likely attributed to the effect of isolation by distance. Other
factors (e.g., small sexual recruitment, somatic mutation and founder effect, etc.) could also play

752 植 物 生 态 学 报 www. plant-ecology.com 32 卷
(e.g., small sexual recruitment, somatic mutation and founder effect, etc.) could also play important
roles in maintaining genetic and clonal diversity in G. racemosa.
Key words genetic diversity, clonal diversity, Globba racemosa, ISSR, mantel test, sexual vs. clonal re-
production
DOI: 10.3773/j.issn.1005-264x.2008.04.003
Many plants use both sexual and clonal re-
productive modes (Richards, 1986). Those plants
may exhibit different degrees of clonal and sexual
reproduction among habitats in response to biotic
and abiotic factors (Barrett, 1980; Abrahamson,
1980; Douglas, 1981), which, in turn, may affect
the genetic structure of populations (Eckert & Bar-
rett 1993; Hangelbroek et al., 2002). In general,
highly clonal populations are expected to exhibit
low genetic diversity because clonal reproduction
yields offspring that are genetically identical to
both the maternal plant and each other (Silander,
1985). However, some studies have demonstrated a
wide range of genetic diversity in populations of
clonal plants or reproduction mode is not always
associated with population genetic structure (Ell-
strand & Roose, 1987; Hamrick & Godt, 1989).
Whether and to what extent reproduction mode can
influence population genetic structure is difficult to
predict.
Population genetic structure is a synthesis pat-
tern resulting from various ecological and genetic
processes or factors, including reproduction, mating
system, habitat conditions, and some random proc-
esses such as the founder effect, mutation and gene
drift (Slatkin, 1987; Avise, 1994; Starfinger &
Stöcklin, 1996; Klimes et al., 1997; Kudoh et al.,
1999). Although it is very difficult to evaluate the
effect of each factor on genetic diversity, significant
correlations were observed between them. For ex-
ample, Loveless and Hamrick (1984) reviewed the
effects of several plant traits (breeding system, flo-
ral morphology, pollination mechanisms, dispersal
modes, life cycles, and population size), and sug-
gested that the breeding system was the principle
factor determining the genetic structure of plant
populations. The effect of reproduction mode, al-
though having received some attention (Hughes &
Richards, 1988; Watkinson & Powell, 1993; Eckert
et al., 2003), has not been clearly determined. Plant
species with a mixed reproductive mode may be an
ideal model to study the function of reproduction on
genetic structure. Investigations linking genetic
structure to reproductive ecology and such eco-
logical factors as habitat differentiation and isola-
tion by distance in species with mixed reproductive
mode may help to understand the processes deter-
mining population genetic structure.
The genus Globba is one of the largest genera
(about 100 species) in the primarily tropical
Zingiberaceae (Kress et al., 2002; Williams et al.,
2004). Globba plants are relatively small (<1 m in
height) understory herbs. There are four species in
Globba in the South and Southwest China: G. ra-
cemosa, G. schomburgkii, G. barthei, and G. lan-
cangensis (Wu, 1981, 1999). These species have
various dominant reproductive strategies: by sexual
seeds (G. lancangensis) or asexual bulbils (G.
schomburgkii and G. barthei). These species repro-
duce predominantly by either sexual seeds or asex-
ual bulbils except G. racemosa which propagates by
seeds or bulbils at different populations.
Codominant markers (allozyme and SSR) are
more ideal in reliable identification of clones (Tor-
res et al., 2003). However, SSR primers are un-
available for most species, and allozyme is not
variable enough in practical research. In recent
years, great efforts have been made to settle this
barrier and many new markers have been devel-
oped. And fortunately, in some cases some domi-
nant markers with enough polymorphism are found
to be as efficient as codominant markers, such as
inter-simple sequence repeat (ISSR) (Zietkiewicz et
al., 1994). SSR offers high polymorphism and is
more stable and reproducible than random amplifi-
cation of polymorphic DNA (RAPD) (Nagaoka &
Ogihara, 1997; Esselman et al., 1999; Qian et al.,
2001; Fernandez et al., 2002). It has been proved to
be very efficient in assessing genetic diversity in
many plant species (Godwin et al., 1997; Joshi et
al., 2000; Camacho & Liston, 2001; Li & Ge, 2001;
Nybom, 2004). Herein, we employed ISSR markers
to assess population genetic and clonal diversity in
G. racemosa.
In the present study, we used ISSR technique
to investigate into the genetic and clonal diversity

4 期 周会平等: 具混合繁殖策略的草本植物异果舞花姜的居群遗传结构 DOI: 10.3773/j.issn.1005-264x.2008.04.003 753
in G. racemosa populations. We analyzed how the
sexual and asexual reproduction variation, isolation
by geographic distance and habitat differentiation
can affect the distribution and level of population
genetic and clonal diversity. Our objective was to
check whether there was a necessary correlation
between reproductive variation and genetic diver-
sity, and whether the correlations between pairwise
genetic and geographic distances among popula-
tions indicated the importance of dispersal traits
and isolation by distance effect. The effects of
founder events and mutation were also touched
upon.
1 Materials and Methods
1.1 The study species
Globba racemosa is an erect herb of 1 (–3) m
in height under the canopy of secondary woods.
The plants flower from June to September and
honeybees are the primary pollinators. Outcrossing
dominates the breeding system. This species regen-
erates through sexual seeds and/or clonal bulbils, as
well as a small number of old rhizomes. Its ellip-
soid capsules, 0.5–1.0 cm in diameter, contain
18–27 seeds. The shuttle bulbils, 0.2–0.5 cm in
diameter, are born on inflorescence. The seeds of G.
racemosa are similar to the seeds of G. lancangen-
sis which ants and mice are the primary seed dis-
persers or predators (Chen et al., 2004). In natural
populations, G. racemosa exhibits high variations in
reproductive complexion among populations (Table
1). In some populations plants produce both nu-
merous bulbils and fruits, while in other popula-
tions, plants produce many bulbils with few or even
no fruits.
Seven G. racemosa populations were found in
Yunnan Province (21°95′–22°55′ N, 99°55′–100°5′
E) (Fig. 1), and great reproduction variation (pre-
dominantly by bulbils or seeds) was observed
among populations. Their vegetative growth (mass
of leaves and stems) is affected by soil nutrients
while reproductive growth (mass of seeds and bul-
bils) is not (Gao et al., 2006). Among the soil fac-
tors, total nitrogen, available nitrogen and organic
compounds contribute greatly to plant growth. The
production of clonal bulbils is significantly corre-
lated to soil fertility while that of sexually produced
seeds is not. Apparently no trade-off exists between
sexual reproduction and asexual reproduction.

Fig. 1 Locations of the seven G. racemosa populations in
Yunnan Province of China


1.2 Genetic and clonal diversity analysis by ISSR
markers
16–20 plants were randomly sampled from
each of the seven populations, and fresh leaves
were collected form these plants. The minimum
distance between sampled plants in each population
was above 10 m. The location of each population
was recorded with GPS, and the distances between
populations were calculated from GPS reads.
Leaves were dried with silica gel and stored in zip-
locked plastic bags until DNA extraction. Genomic
DNA was extracted using a modified cetylmethyl-
ammonium bromide (CTAB) method of Doyle and
Doyle (1987).
15 ISSR primers (UBC primer set No. 9, Bio-
technology Laboratory, University of British Co-
lumbia, British Columbia, Canada) were used in
this study, which all produced bright and repro-
ducible bands (Table 2). Polymerase chain reaction
(PCR) amplification was undertaken in a total reac-
tion volume of 20 μl containing 2 μl of 10 × PCR
buffer (100 mmol·L-1 Tris-HCl [pH 8.3], 500
mmol·L-1 KCl, 15 mmol·L-1 MgCl2), 100 μmol·L-1
dNTP, 0.375 μmol·L-1 primer, 50 ng DNA template,
and 1.5 U Taq polymerases (TaKaRa Biotech Inc.,
Dalian, China). The reaction was performed in an
Eppendorf master cycler, and commenced with an
initial denaturation step at 95 ºC for 3 min, fol-
lowed by 35 cycles of 25 s at 94 ºC, 30 s at anneal-
ing temperature (Table 2), 1.5 min at 72 ºC, and a
final 7 min extension at 72 ºC. Amplification prod-
ucts were resolved on 1.5% agarose gels stained
with ethidium bromide (EB). After electrophoresis,
the gels were visualized and photographed using the

754 植 物 生 态 学 报 www. plant-ecology.com 32 卷

Table 1 Geographical, physical habitat and reproductive characteristics of seven Globba racemosa populations (Data of
Pop2–Pop7 are from Gao et al., 2006)
Pop1 Pop2 Pop3 Pop4 Pop5 Pop6 Pop7
Latitude/longitude

21º53′ N/
100º58′ E
22º53′ N/
99º48′ E
24º04′ N/
97º48′ E
24º05′ N/
99º44′ E
24º25′ N/
97º50′ E
24º35′ N/
97º40′ E
24º56′ N/
102º29′ E
Habitat size (100 m2) 5–10 5–10 10–20 5–10 10–20 10–20 10–20
Altitude (m) 1 350 1 700 1 136 1 873 1 456 1 352 2 100
Soil water supply Rich Rich Rich Poor Very rich Poor Very poor
Degree of disturbance Slight Slight Undisturbed Intense Undisturbed Intense Slight
No. of fruits per plant
(Mean ± 1SE, n = 30)
1.65 ± 0.37 6.53 ± 0.62 0 0.80 ± 0.228 19.10 ± 1.58 0 0.20 ± 0.12
No. of bulbils per plant
(Mean ± 1SE, n = 30)
20. 12 ± 0.88 24. 67 ± 1.88 30.00 ± 1.90 24.27 ± 1.79 28.63 ± 0.77 18.83 ± 1.28 24.43 ± 1.30
Ratio of fruits/bulbils
(Mean ± 1SE, n = 30)
0.11 ± 0.03 0.34 ± 0.06 0 0.04 ± 0.01 0.67 ± 0.05 0 0.01 ± 0.01


Table 2 Fifteen ISSR primers used for Globba racemosa
amplification
Primer
No.
Sequence Annealing temperature
(ºC)
807
808
810
835
836
840
847
848
855
857
886
887
888
889
890
(AG) 8 T
(AG) 8 C
(GA) 8 T
(AG) 8 YC
(AG) 8 YA
(GA) 8 YT
(CA) 8 RC
(AG) 8 RG
(AC) 8 YT
(AC) 8 YG
VDV (TC) 7
DVD (TC) 7
BDB (CA) 7
BDB (AC) 7
VHV (GT) 7
50
52
50
50
52
50
52
52
52
52
52
52
52
52
52
B=C/G/T, D=A/G/T, H=A/C/T, R=A/T, V=A/C/G,
Y=C/T


GeneSnap Bio Imaging System (Syngene, Freder-
ick, MD, USA). Molecular weights were estimated
using a 100 bp DNA ladder (Sangon Inc., Shanghai,
China).
1.3 Data analysis
ISSR bands were scored as present (1) or ab-
sent (0) for each DNA sample and presented in a
binary data matrix. Bands were analyzed to inves-
tigate the overall distribution of genetic variance
using Analysis of Molecular Variance (AMOVA)
software (Excoffier, 1993) under Hardy-Weinberg
equilibrium. Three population genetic parameters
calculated from POPGENE (Yeh et al., 1997) were
adopted to assess the levels of population genetic
diversity: Nei’s gene diversity (h), Shannon’s in-
formation index (I) and percentage of polymorphic
loci (PPB).
Cluster analysis using the unweighted
pair-group method with arithmetic averages (UP-
GMA) (Sneath & Sokal, 1973) by NTSYS (Rohlf,
1997) was used to generate a dendrogram of all 130
individuals based on Nei’s genetic distance (Nei,
1972). The clonal diversity of each population was
calculated as G/N, where G is the number of geno-
types and N is the total number of individuals ana-
lyzed (Wolfe & Randle, 2001). G/N has a range of 0
to 1, where 0 indicates that all samples in a popula-
tion have the same multilocus genotype, and 1 in-
dicates each sample in a population has a unique
multilocus genotype. The UPGMA dendrogram of
all 130 samples from seven populations was con-
structed using Nei’s genetic distance with POP-
GENE. Pairwise genetic distances among popula-
tions were also shown.
Evidence of isolation by distance among
populations was obtained by examining the correla-
tion between matrices of pairwise genetic distances
and geographical distances (Slatkin, 1993). Sig-
nificance of the observed relationships was ob-
tained using a Mantel test (Mantel, 1967).
Reproductive variation is reflected by number
of fruits, bulbils, and ratio of fruits to bulbils. Lev-
els of within-population genetic diversity are indi-
cated by h, I and PPB. The Pearson correlation
function from SPSS software was adopted to esti-
mate the relationships between genetic diversity
and reproduction variation.
2 Results
Fifteen primers produced a total of 118 clear
and reproducible amplification fragments (7.9
bands per primer on average) over the 130 plant

4 期 周会平等: 具混合繁殖策略的草本植物异果舞花姜的居群遗传结构 DOI: 10.3773/j.issn.1005-264x.2008.04.003 755
individuals, ranging in size from 300 to 1 900 bp.
84 out of the 118 loci were polymorphic (PPB =
71.19%). PPB values varied from 5.08% to 33.90%
among populations.
AMOVA analysis indicated a large proportion
of genetic variation (59.44%) resided among popu-
lations rather than within populations. The result is
consistent with the GST value (0.590 7) from POP-
GENE. Pop 6, Pop 4 and Pop 5 showed relatively
high genetic diversity while Pop 7 had the lowest
variation (Table 3).


Table 3 Genetic variability within populations of Globba racemosa detected by ISSR analysis
Population Sample size na ne h I PPB (%)
Pop1
Pop2
Pop3
Pop4
Pop5
Pop6
Pop7
Average
Species level
20
19
16
17
19
19
20
18.57
130
1.245 8
1.254 2
1.245 8
1.330 5
1.305 1
1.339 0
1.050 8
1.253 0
1.711 9
1.141 2
1.181 3
1.206 4
1.201 5
1.228 3
1.222 9
1.025 7
1.172 5
1.398 7
0.083 7
0.099 2
0.109 5
0.117 9
0.125 5
0.125 3
0.014 7
0.096 5
0.236 3
0.125 7
0.143 8
0.155 8
0.176 4
0.181 3
0.184 5
0.022 3
0.127 1
0.357 5
24.58
25.42
24.58
33.05
30.51
33.90
5.08
25.30
71.19
na: number of alleles ne: effective number of alleles h: Nei’s gene diversity I: Shannon’s information
index PPB: percentage of polymorphic loci


The dendrogram of 130 individuals showed that
all individuals were allocated into 116 ISSR geno-
types according to the 118 ISSR markers. All indi-
viduals from one population formed a clade before
clustering together with other such clades, and none
of the 113 genotypes were shared by more than one
population. Five populations (Pop 1, 2, 4, 5 and 6)
had individual unique genotypes (G/N = 1.0), whereas
Pop 7 had a much lower clonal diversity (G/N = 0.45).
The mean G/N value of this species was 0.88.
The dendrogram of seven populations pro-
duced three large clusters: Pop 2 and Pop 4; Pop 3,
Pop 5, Pop 6 and Pop 7; Pop 1 (Fig. 2). The Mantel
test indicated that genetic distances among popula-
tions were significantly correlated with geographi-
cal distances (r = 0.68, p = 0.012 < 0.05) (Fig. 3).
Field investigation showed slight difference in
bulbil production and great variation in seed output
among those study populations. However, no sig-
nificant correlation (p > 0.05) was found between
population genetic diversity (PPB, I and h values)
and sexual vs. clonal reproduction level (number of
fruits, number of bulbils and ratio of fruits to bul-
bils).
3 Discussion
Our field investigation showed that there was
slight difference in bulbil production and great
variation in seed output among the seven study
populations. The variation in propagule output
probably resulted from the difference of soil condi-
tions, local weather and available pollinators due to
environmental landform and elevation differentia-
tion (Ceplitis, 2001; Young et al., 2002; Gao et al.,
2006). In the present study, seven populations of G.
racemosa showed different and relatively high
within-population genetic and clonal variation.
Most of the genetic variation (ΦST = 0.594 4) ex-
isted among populations.



Fig. 2 UPGMA dendrogram based on Nei’s coefficients
illustrating the genetic relationship
among seven studied populations


For those species with mixed reproductive
mode, the production of different types of
propagules is expected to be associated with dif-
ferent levels of genetic and clonal diversity
(Hughes & Richards, 1988; Ronsheim & Bever,

756 植 物 生 态 学 报 www. plant-ecology.com 32 卷
2000; Eckert et al., 2003). Theoretically, predomi-
nantly clonal species should exhibit lower
within-population genetic diversity than predomi-
nantly sexual species because of the absence of
gene recombination and high spread capacity al-
though there are some exceptions (Silander, 1985;
Ashton & Mitchell, 1989; Piquot et al., 1998).
However, several reviews showed that clonal plants
may also have high genetic diversity (Ellstrand &
Roose, 1987; Widén et al., 1994). In our study, al-
though sexual fruit set and within-population ge-
netic diversity varied greatly among populations,
neither the number of fruits and bulbils, nor their
ratio was significantly correlated to within- popula-
tion genetic diversity. Most G. racemosa plants
produced many asexual bulbils, but asexual repro-
duction did not seem to reduce within-population
genetic diversity. Our results suggest that variation
in levels of asexual vs. asexual reproduction is not a
major determinant of the genetic differentiation
among those populations. Other factors, such as
isolation by distance, genotype-environment inter-
actions, somatic mutation, founder effect and other
random processes may play important roles in gen-
erating high within-population genetic diversity.



Fig. 3 Correlation analysis of genetic distances and geo-
graphical distances among populations by
Mantel test (r = 0.68, p = 0.012).



We found a significant correlation between
genetic and geographical distance among popula-
tions (Fig. 3). In general, gene flow decreases with
the increase of distance between populations, and
consequently the differentiation increases (Hamrick
& Godt, 1989). Our result indicates an effect of
isolation by distance, and may suggest that limited
gene migration greatly contributes to the differen-
tiation among G. racemosa populations. G. ra-
cemosa flowers are primarily pollinated by honey-
bees, which do not cover large distances during
foraging, particularly over fragmental areas (pers.
obs.). The small seeds are likely removed by ants
which have limited dispersal ability or home range.
Therefore, G. racemosa is likely to have very short
seed dispersal distance. Additionally, many popula-
tions use clonal reproduction as the main propaga-
tion mechanism, and clonal bulbils are considered
to be more disadvantageous for long-distance dis-
persal than seeds for their susceptibility during long
migration process. Therefore, successful gene mi-
gration through pollen or propagules probably oc-
curs only locally or between adjacent populations,
and will be reduced with the increase of geo-
graphical distance.
In this study on G. racemosa, the mean value
of clonal diversity was 0.88, which is much higher
than the averages reported for other clonal plants
(G/N = 0.43, Jacquemyn et al. (2005); G/N = 0.17,
Ellstrand & Roose (1987)). In G. racemosa, there
was no significant correlation between clonal di-
versity and clonal reproduction levels. This result is
congruent with the claims of Hamrick and Godt
(1989) and Eckert et al. (2003) that there is no
necessary association between clonal diversity and
reproductive modes.
The extremely high clonal diversity in G. ra-
cemosa is likely resulted from other factors other
than reproductive mode, since clonal diversity is
impressible to many drivers such as mating system,
mutation, environment, and founder effects. Be-
cause phenotypic traits influence plant fitness, pat-
terns of differential survival and reproduction
unique to each microsite could lead to the evolution
of localized clonal structure (Schemske, 1984). As
pointed out by Welch and Meselson (2000, 2001),
in a purely clonal diploid population, we may ex-
pect that new mutations accumulate in different
clones at both homologous alleles and we may then
expect a higher conservation of allele number with
clonality (this is the so-called “Meselson effect”).
In addition, Watkinson and Powell (1993) showed
that the input of only a few seedlings each year into
a population of ramets accelerated the loss of foun-
der genets but acted as a powerful mechanism for
maintaining clonal diversity in populations.
Furthermore, the higher the incomparability is, the

4 期 周会平等: 具混合繁殖策略的草本植物异果舞花姜的居群遗传结构 DOI: 10.3773/j.issn.1005-264x.2008.04.003 757
greater the contribution of sexual reproduction to
clonal diversity. The high clonal diversity in G. ra-
cemosa may indicate that forces such as phenotypic
fitness, sexuality introducing, self-incompatibility
and mutation tend to promote population diver-
gence.
Founder effect is also an important factor in-
fluencing clonal diversity (Baker & Moeed, 1987),
which will decrease with time and counteracted by
subsequent forces after founder event (Watkinson &
Powell, 1993; Houliston & Chapman, 2004). Pop 7
might provide an evidence for a severe founder ef-
fect in G. racemosa. This population had much
lower clonal diversity than other populations, al-
though there were some seeds produced. Taking
into account the long distance of Pop7 away from
other populations (Fig. 1), we suspect it is a newly
established population with certain genets through
an unknown dispersal process. Although there are
some new seedlings of sexually produced seeds and
mutation is happening, they can not overcome the
influence of founder genets yet. Founder event in
Pop7 still plays a dominant role in maintaining
clonal diversity of this population at present.
Globba species are distributed throughout
tropical and parts of subtropical Asia, ranging from
India to southern China, south and east to Philip-
pines and New Guinea (Schumann, 1904), with the
center of distribution in monsoonal Southeast Asia,
especially Thailand (Larsen, 1996) and Myanmar
(Kress et al., 2003). In our study, the geographical
distribution coincides with the genetic relationships
of the seven studied populations. This may suggest
a northward spread tendency of G. racemosa in
China (Fig.1, Fig.2). Therefore, we guess, as the
northern boundary of world Globba, the G. ra-
cemosa in Yunnan could originate southerly from
the distribution center in Thailand or Myanmar
which spread northerly to China. This inference
still needs the validation of phylogenetic and bio-
geographic studies in the future.
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