全 文 ：尖叶拟船叶藓营养元素生殖配置格局
?
李 菁 , 魏 华 , 刘 冰 , 王亚琴 , 陈 军
(吉首大学植物资源保护与利用湖南省重点高校实验室 , 湖南 吉首 416000)
摘要 : 为了揭示尖叶拟船叶藓营养元素生殖配置规律 , 本文对其 12 种营养元素的生殖配置格局和季节动
态进行了研究。其结果如下 : 成熟孢子体的生物量配置为 6 .67% ; 在成熟孢子体中 12 种营养元素的含量
顺序是 : C (452 mg?g) > N ( 35 mg?g) > K ( 8439.9μg?g) > Ca ( 7012.9μg?g) > P ( 2129 .2 μg?g) > Mg
( 1482 .9μg?g) > Na ( 432 .9μg?g) > Mn ( 196 .3μg?g) > Fe (177 .7μg?g) > Al (174 .8μg?g) > Zn (68.1μg?g)
> Cu (19.4μg?g) ; 成熟孢子体中营养元素生殖配置顺序是 : K (17 .7 % ) > P (15 .1% ) > Cu ( 13 .3% ) > N
( 11 .6% ) > Na ( 10 .5 % ) > Mn ( 7. 8% ) > Zn ( 7. 5 % ) > C ( 6. 9 % ) > Mg ( 6 .8 % ) > Ca (5 . 4% ) > Fe
( 1. 3 % ) > Al (1 . 2% )。
关键词 : 苔藓植物 ; 营养元素 ; 生殖配置格局
中图分类号 : Q 948.1 文献标识码 : A 文章编号 : 0253 - 2700 (2009) 03 - 219 - 08
Study on the Reproductive Allocation Pattern of
Nutrients in Dolichomitriopsis diversiformis
(Bryopsida: Lembophyllaceae)
LI J ing, WEI Hua, LIU Bing, WANG Ya-Qin, CHEN Jun
( Key Laboratory of Plant Resources Conservation and Utilization, J ishou University, J ishou 416000 , China)
Abstract : In order to reveal the reproductive patterns of Dolichomitriopsis diversiformis, twelve nutrients allocation patterns
and seasonal dynamics were studied . The results were as follows: mature sporophyte biomass allocation was 6 .67% ; 12
nutrient contents in mature sporophytefollowed the order: C ( 452 mg?g) > N (35 mg?g) > K ( 8439 .9μg?g) > Ca
( 7012 .9μg?g) > P (2129 .2μg?g) > Mg (1482 .9μg?g) > Na (432 .9μg?g) > Mn (196 .3μg?g) > Fe (177 .7μg?
g) > Al (174 .8μg?g) > Zn (68 .1μg?g) > Cu (19.4μg?g) ; nutrient allocation patterns inmature sporophyte followed
the order: K (17 .7% ) > P (15 .1% ) > Cu ( 13.3 % ) > N (11 .6% ) > Na (10 .5% ) > Mn (7 . 8% ) > Zn
( 7. 5 % ) > C (6 . 9% ) > Mg (6 . 8% ) > Ca (5. 4 % ) > Fe (1. 3 % ) > Al (1 . 2% ) .
Key words: Bryophyte; Nutrients; Reproductive allocation pattern
Reproductive allocationof resources is defined as
the proportion of resources allocated to the reproduc-
tive parts relative to the total nutrients in the reproduc-
tive process of a plant ( J iang, 1992 ) . Plants act in
such a way that shortage or excess of a resource will
influence the way in which other resources are ac-
quired and allocated, and that the resulting adjustment
optimizes plant performance . Keeping a balance be-
tween resources entails homeostatic adjustment of re-
source concentrations by alteration of resource alloca-
tion patterns (Chapin et al. , 1987) .
In bryophytes, it appears that spore production
(i . e ., sexual reproduction) comes at a great energy
cost including assumption or reallocation of nutrients .
云 南 植 物 研 究 2009 , 31 (3) : 219～226
Acta Botanica Yunnanica DOI : 10 .3724?SP. J . 1143 .2009.08074
? ?Foundation item: The National Natural Science Foundation of China ( Grant No . 30470181)
Received date: 2008 - 12 - 13 , Accepted date: 2009 - 03 - 04
作者简介 : 李菁 ( 1955 - ) 男 , 教授 , 主要从事植物生态学和保护生物学研究。E-mail: lkg@ jsu. edu. cn
Bisang and Ehrlén ( 2002 ) proved that the gameto-
phyte and sporophyte must compete for limited re-
sources ( like nutrients) within the plant, which may,
to some extent, contribute to the rarity of perigonium
and sporophyte in some bryophyte species because of
the high reproductive costs (Stark et al. , 2000 ) .
Dolichomitriopsis diversiformis ( Mitt .) Nog .,
which belongs to Lembophyllaceae, is one dioecious
moss species . The morphology is remarkably different
between its perigonium and perichaetium, the number
of perichaetia is much more than that of perigonia,
and the perigonia are very rare and difficult to find
(Liu et al. , 2006a) . It was found that the frequency
of sporophyte abortion in this moss was 7 .3% ( Li et
al. , 2007) . For a better understanding of the mecha-
nisms underlying the sporophyte development and the
life cycle of this moss, it is essential that we examine
the reproductive allocation and seasonal dynamics of
nutrients in reproductiveprocess of thisplant . Howev-
er, to our knowledge, very fewstudies have addressed
the reproductive allocation and seasonal dynamics of
nutrients in bryophytespecieswith respect to thesexu-
al reproduction, which may due partly to the difficul-
ties in determining gametophytebiomass and in identi-
fying parts serving reproduction ( Bisang and Ehrlén,
2002) .
The objectives of this study were to investigate
the nutrient allocation patterns, to compare the sea-
sonal dynamics in 12 nutrient contents (C , N, P , K ,
Ca, Na, Mg, Fe, Zn, Mn, Al and Cu) in reproduc-
tive and vegetative parts, and to discuss whether the
sporophyte development depend on the availability of
certain nutrients in the moss D. diversiformis . Most
studies on reproductive allocation patterns were per-
formed on tracheophyte and only a few on bryophytes,
hence, the study on nutrient allocation patterns in
moss will assist further research on the reproductive
allocation of bryophyte species .
1 Materials and methods
1 .1 Site description
The studywasconducted in FanjingMountain (27°49′50″-
28°11′30″N, 108°45′55 ″- 108°48′30 ″E) in the eastern part of
Guizhou, China (Liu et al. , 2006a) . The annual precipitation
ranges from1100 to2600 mm, the annual mean relativehumidi-
ty is above80% , and the annual temperature ranges from 5 to
17℃ (Li et al. , 2002 ) . The moss D. diversiformis is one of
theendemic bryophytes inEast Asiawith limiteddistributionar-
eas (Wu, 1992) . In Fanjing Mountain, the species are mainly
distributed in the forest community which is dominated by Cy-
clobalanopsis stewardiana and Sinarundinaria nitidacommunit
ranges from1650 mto 2080 m elevation (Liu et al. , 2006b) .
1 . 2 Species studied
D. diversiformis is a perennial dioecious species, which
mainlygrows on tree trunks ( epiphytic) , and are seldom found
on rocks . The perigonium is rarely found, and the development
of the plant population dependslargelyonvegetativepropagation
rather than sexual reproduction . Stark ( 2002 ) suggested 20
stages for scoringthe developmental stages of bryophyte species,
centering on the reproductive phases only . As for the rarity of
sporophyte andthe difficulties infinding perigonium, we scoring
the developmental stages of this moss simply as five important
stages based on a long period of field observations (fromApril,
2004 to January, 2006) :
GG: gametophyte rapid growing stage;
PG: perichaetiumgrowing stage, i .e ., before fertilization;
SI : ?sporophyte developing initial stage . This corresponds
with the development of the embryofollowingfertiliza-
tion;
YS: ?young sporophyte growing stage (capsule is green with
calyptra on it) ;
MS: ?mature sporophyte stage (capsule is brown and green
with an intact calyptra) .
1 . 3 Sampling and measurements
Plant materials from certain populations were collected on
February, April , July, September, November and December,
2006 (Table 1 ) , respectively . Specimens ( B . Liu 05 - 102 )
were deposited at the Plant Herbarium of Jishou University
( JIU) . The sampling sites were in theelevation about 1900 m .
The choice of the sites was subjective because most sporophyte
plants were found near this elevation . All sampleswere collect-
edwithin a short period without precipitation . In the reproduc-
tive period ( fromSeptember to December, 2006 ) , only plants
bearing sporophyteswere sampled . Plant materials collected ( -
60 g) were rinsed several times with distilled water after remov-
ing of old brown parts .
The reproductive parts ( only include perichaetium and
sporophyte because the perigonium can hardly be found) were
carefully separated from the vegetative parts ( gametophyte) un-
der stereo microscope, then they were oven-dried to a constant
weight at 80℃ for 24 h and weighed to the nearest 0 .0001 g,
separately . After ground to finepowderwith an electric grinder,
022 云 南 植 物 研 究 31 卷
the samples were placed in paper bags, and stored frozen up
until nutrients analyses . Before analyses for nutrients the sam-
pleswere oven-dried to aconstantweight again (80℃ for 24h) .
After digesting, total C and N contents were determined on an
elementary analyzer ( Vario EL III , made in German) . Total
nutrient contents (P , K , Ca, Na, Mg, Fe, Zn, Mn, Al and
Cu) were determined by the plasma atomic emission spectrome-
ter ( IRIS Intrepid II XSP , madein America) . All theanalyses
were performed at the Center of Analysis and Test of Wuhan
University . The laboratory included its own blank and standard
samples . The nutrient allocation (NA) to a plant part was esti-
mated as follows:
NA = mnutrient mass of reproductive parts?total nutrient mass of
the plant
= nutrient content of reproductive parts×corresponding pro-
portion of dry mass?( nutrient content of reproductive parts×
corresponding proportion of dry mass+ nutrient content of vege-
tative parts×corresponding proportion of dry mass)
2 Results and analysis
2 .1 Biomass allocation
As was shown in Table 1 , biomass allocation to
reproductive parts increased with the development of
sporophyte, it was only 0 .74% at PG stage, but in-
creased morethan three times at SI stage, and reached
its peak of 6 .67% at MS stage .
2 . 2 Nutrients allocation patterns
Reproductive allocation patterns and seasonal dy-
namics of nutrients in themoss D. diversiformis at dif-
ferent reproductive stages were shown in Table 2 and
Fig . 1 .
The highest content of carbon ( C ) was 452 mg?
g . No significant difference was found in C content
between vegetative and reproductive parts, and the
content of C was very stable at different stages .
Nitrogen (N) content showed a similar trend in
both vegetative and reproductive parts during PG and
YS stage, it ranged from 19 to 26 mg?g in vegetative
parts, while in mature sporophyte the content was 35
mg?g which was 1 .8 times as high as vegetative
parts .
A constant trend was found in phosphor (P) ac-
cumulation in the reproductive parts, with the highest
content of 2129 .2μg?g which was about 2 .5 times as
high as that of the vegetative parts at MS stage . Dur-
ing SI and MS stages, P contents were always much
higher in reproductive parts than in vegetative parts .
In vegetative parts, P content dropped with the devel-
opment of sporophyte (from 1114 .2 to 855 .8μg?g) .
Kalium (K ) accumulative pattern appeared to be
similar to thatof P . K alsoaccumulatedsteadily in re-
productive parts with the development of sporophyte,
with the highest content of 8439 .9μg?g which was 3
times as high as that of vegetative parts at MS stage .
No significant differenceof K content was found in ve-
getative parts at different stages .
Calcium (Ca) content was 9212 .4μg?g in vege-
tative parts at PG stage coupled with the highest Ca
content (12673.4μg?g) inperichaetium . It decreased
by 44 .7% in mature sporophyte ( 7012 .9μg?g) while
did not vary strongly in vegetative parts .
Natrium (Na) contents in reproductivepartswere
on average29 .2% higher than invegetative parts after
PG stage . Na contents did not vary strongly invegeta-
tive parts .
When perichaetia began to develop, magnesium
(Mg) content increased on average by 46 .6% inveg-
etative parts . The highest Mg content ( 1662 .4μg?g)
in vegetative parts was coupled with the highest Mg
content ( 2254 .7μg?g) in perichaetium at PG stage .
After a slightly decrease during SI and YS stage, Mg
content increased again at MS stage .
Table 1 Biomass allocation to vegetative and reproductive parts in the moss D. diversiformis
Developmental
stages
Sampling date
Biomass allocation ( % )
Gametophyte Perichaetium Sporophyte
GG 17th, April 100 ?. 00 — —
PG 27 ?th, J uly 99 . 26 0 . 74 —
SI 30 ?th, September 97 . 07 — 2 . 93
YS 1 ?st, November 95 . 23 — 4 . 77
MS 10 ?th, December 93 . 33 — 6 . 67
Notes: The“—”indicates that there is no reproductive parts exist .
1223 期 LI Jing et al. : Studyon the Reproductive Allocation Pattern of Nutrients in Dolichomitriopsis diversiformis . . .
Table 2 Reproductive allocation patterns and seasonal dynamics of nutrients in the moss D. diversiformis at different reproductive stages
Nutrient Developmental stage
Structure content
Gametophyte Perichaetium Sporophyte
NA ( % )
GG 439 M— — 0 ?
PG 435 M429 ?— 0 v. 7
C SI 434 M— 422 ?2 v. 9
YS 437 M— 429 ?4 v. 7
MS 433 M— 452 ?6 v. 9
GG 19 ;— — 0 ?
PG 252 M3 \— 0 v. 7
N SI 26 ;— 28 ?3 v. 1
YS 19 ;— 22 ?5 v. 5
MS 19 ;— 35 ?11 .6
GG 800 (. 7 — — 0 ?
PG 1114 ;. 2 838 \. 6 — 0 v. 6
P SI 777 (. 6 — 1512 ?. 9 5 v. 5
YS 1009 ;. 2 — 1862 ?. 5 8 v. 5
MS 855 (. 8 — 2129 ?. 2 15 .1
GG 2615 ;. 1 — — 0 ?
PG 2735 ;. 8 2708 o. 7 — 0 v. 7
K SI 2125 ;. 7 — 4447 ?. 5 5 v. 9
YS 2524 ;. 7 — 6966 ?. 9 12 .1
MS 2799 ;. 4 — 8439 ?. 9 17 .7
GG 7982 ;. 9 — — 0 ?
PG 9212 ;. 4 12673 ?. 4 — 1
Ca SI 7409 ;. 6 — 7138 ?. 6 2 v. 8
YS 9064 ;. 2 — 6599 ?. 3 3 v. 5
MS 8746 ;. 1 — 7012 ?. 9 5 v. 4
GG 219 (. 7 — — 0 ?
PG 307 (. 1 472 \. 8 — 1 v. 1
Na SI 272 (. 6 — 480 ?. 2 4
YS 290 (. 8 — 388 ?. 6 6 v. 3
MS 264 (. 5 — 432 ?. 9 10 .5
GG 1014 ;. 2 — — 0 ?
PG 1662 ;. 4 2254 o. 7 — 1 ?
Mg SI 1461 ;. 1 — 1647 ?. 5 3 v. 3
YS 1363 ;. 3 — 1174 ?. 3 4 v. 1
MS 1461 ;. 2 — 1482 ?. 9 6 v. 8
GG 865 (. 1 — — 0 ?
PG 1530 ;. 5 807 \. 1 — 0 v. 4
Fe SI 2023 ;. 1 — 160 ?. 4 0 v. 2
YS 1715 ;. 6 — 260 ?. 3 0 v. 8
MS 972 (. 6 — 177 ?. 7 1 v. 3
GG 80 ?. 0 — — 0
PG 62 ?. 8 170 \. 2 — 2
Zn SI 93 ?. 5 — 125 .7 3 v. 9
YS 79 ?. 3 — 81 ~. 9 4 v. 9
MS 60 ?. 3 — 68 ~. 1 7 v. 5
GG 151 (. 0 — — 0 ?
PG 155 (. 3 150 \. 1 — 0 v. 7
Mn SI 322 (. 8 — 425 ?. 7 3 v. 8
YS 247 (. 7 — 258 ?. 1 5
MS 165 (. 4 — 196 ?. 3 7 v. 8
GG 738 (. 2 — — 0 ?
PG 1438 ;. 1 843 \. 6 — 0 v. 4
Al SI 1882 ;. 9 — 161 ?. 4 0 v. 3
YS 1904 ;. 1 — 285 ?. 3 0 v. 7
MS 1006 ;. 3 — 174 ?. 8 1 v. 2
GG 12 ?. 3 — — 0
PG 13 ?. 2 21 J. 4 — 1 v. 2
Cu SI 14 ?. 9 — 11 ~. 8 2 v. 3
YS 11 ?. 9 — 17 ~. 6 6 v. 9
MS 9 ?. 0 — 19 ~. 4 13 .3
Note: The units of all the nutrient contents areμg?g, except C and N ( mg?g) ; the“—”indicates that there are no reproductive parts exist
222 云 南 植 物 研 究 31 卷
Fig . 1 a- l : Seasonal dynamics of 12 nutrients in vegetative ( gametophyte) and reproductive parts ( perichaetium or sporophyte)
in the moss D. diversiformis . Here, GG, PG, SI , YS, and MS represent different developmental stages
3223 期 LI Jing et al. : Studyon the Reproductive Allocation Pattern of Nutrients in Dolichomitriopsis diversiformis . . .
Ferrum ( Fe) contents were relatively higher in
vegetative parts than in reproductive parts . In vegeta-
tive parts, Fe content increased gradually before SI
stage, with the highest content of 2023 .1μg?g, and
then it decreased with the development of sporophyte .
Fe content in reproductive parts decreased sharply after
fertilization ( i . e ., after PG stage) , from 807.1 to
160 .4μg?g .
At PG stage, zinc (Zn) content was 2 .7 times as
high in reproductive parts as in vegetative parts . Zn
content declined steadily with the development of
sporophyte, with the lowest content ( 68 .1 μg?g) in
mature sporophyte .
Surprisingly, manganese (Mn) content increased
sharply in both vegetative and reproductive parts at SI
stage, which wereover 2 times as high as thatof previ-
ous period, and then both decreasedgraduallywith the
development of sporophyte .
Aluminium (Al) contents were relatively higher in
vegetative parts than in reproductive parts . In vegeta-
tive parts, Al increased rapidly before YS stage, with
the highest content of 1904 .1μg?g, and then decreased
sharply when the sporophyte became mature ( from
1904.1 to 1006 .3μg?g) . Al content in reproductive
parts decreased sharply after fertilization ( from 843 .6
to 161 .4μg?g) .
The highest copper ( Cu) content ( 21 .4 μg?g)
was found inperichaetiumat PG stage, after adecrease
at SI stage it increased again with the development of
sporophyte . Cu content in vegetative parts increased
slightly beforeSI stage, and thendecreased (from14 .9
to 9 .0μg?g) .
3 Discussion
In comparison withother bryophyte species, D. -
diversiformis had a relatively lower biomass allocation to
mature sporophyte ( 6 . 67% ) . For example, the bio-
mass allocation to mature sporophyte was 16% in Di-
cranumpolysetum (Ehrlén et al. , 2000) , and 30% in
Tetraphis pellucid ( Kimmerer, 1991 ) . This indicates
that D. diversiformis, as a perennial species, allocates
much lower resources to reproductive parts than that of
vegetative parts and is largely depends on vegetative
propagation for population development .
Nutrient contents in mature sporophyte followed
the order: C ( 452 mg?g) > N ( 35 mg?g) > K
(8439 .9μg?g) > Ca (7012 .9μg?g) > P ( 2129 .2
μg?g) > Mg ( 1482 .9μg?g) > Na ( 432 .9μg?g) >
Mn (196 .3μg?g) > Fe (177 .7μg?g) > Al ( 174 .8
μg?g) > Zn (68 .1μg?g) > Cu (19 .4μg?g) .
C made up a very large proportion of plant dry
mass which was high above the contents of other nutri-
ents . N and P are essential in making proteins and
DNA , and P is needed in ATP to maintain energy . K
is mainly located in the most active parts of a plant
which is a translocatable nutrient that can move easily
to theyoung parts of a plant . High contents of N, P,
and K coincided with the capacity for rapid growth in
reproductive conditions of D. diversiformis . The rapid
contents accumulation in developing sporophyte can be
explained as: N, P andK aremoved fromvegetativeto
reproductive parts, or reproductive parts actively up-
take these three nutrients (Gerdol et al. , 2005; Rydin
and Clymo, 1989 ) .
In this paper, as there was no significant differ-
ence in K contents in vegetative parts at different stag-
es, the strong increase of K could be explained mainly
as the active uptake of K by developing sporophyte .
While the increase of N and P could be explained as
both the translocation of N and P from gametophyte to
sporophyte and the active uptakeof N and P by devel-
oping sporophyte .
As is known Na is beneficial to the growth of C3
plant (Pan, 2004) , the high Na content was found in
the rapid growing sporophyte in the moss D. div-
ersiformis, indicating that Namight be beneficial to the
growth of this moss . Raven et al . ( 1998 ) have re-
viewed the evidence for the C3 pathway in bryophytes,
and our result indirectly corresponds with such evi-
dence .
The decrease of Ca and Zn contents in reproduc-
tive parts may partly due to the growth of sporophyte
which“diluted”their contents, and the stability of Ca
content in vegetative parts could be explained as the
immobile feature of Ca .
Mg is one of the major constitutes of chlorophyll ,
422 云 南 植 物 研 究 31 卷
hence the pattern of seasonal changes in Mg content in
different parts of the moss suggest different photosyn-
thetic activity . During reproductive stages, both the
gametophyte and sporophyteseemed to havehighphoto-
synthetic activity . The highest content of Mg in
perichaetium indicates that it had a high photosynthetic
activity .
Fig. 1h and Fig. 1k showed obvious differences in
Fe and Al contents between vegetative and reproductive
parts . One of the factors contributing to high contents
of Fe and Al ingametophyte is that as cells becomeold
or other ions move to thesporophyte, new binding sites
are exposed, permitting moreof these ions to accumu-
late there, and may also partly due to the rapidgrowth
of sporophyte .
Cu is one of constitutes of plastocyanin in chloro-
plast, and play a role in photosynthesis . Therefore,
high Cu content insporophytesuggestted ahigh capaci-
ty of photosynthesis and a high demand for Cu ( Pan,
2004) .
Mn is an activator of several enzymes, such as
dehydrogenase, decarboxylase, kinase, oxidase and so
on, and especially has an effect on Tricarboxylic Acid
Cycle ( Pan, 2004 ) . So, the high Mn content in
perichaetium suggested a high rate of biosynthesis in
thedeveloping sporophyte and also a high demand for
Mn after fertilization .
In this study, the results that all contents of the
12 nutrients inmature sporophyteweregreater than that
of gametophyte at MS stage except Ca, Fe and Al part-
ly supports the assumption that sporophyte development
is the most expensive developmental stage in terms of
demands for resources ( Rydgren, 2003 ) , indirectly in-
dicating that the developing sporophyte of compete nu-
trients with gametophyte in themoss D. diversiformis .
Mosses mostly depend upon atmospheric deposi-
tion for their supplies of water and mineral nutrients
(Schintu et al. , 2005 ) which was especially the case
for the epiphytic mosses . Research on substrate-related
element uptakeby epiphytic lichens has shown minimal
accumulation due to what is probably an overall defi-
ciency of elements inbark tissues (DeBruin and Hack-
enitz, 1986 ) . Also, bryophytes seem to have uptake
specificity for things they need over things they do not .
For example, the thallose liverwort Dumortiera hirsuta
preferentially took up Ca, Mg, and Znover Cd (Maut-
soe and Beckett, 1996) .
In this paper, N, P, K , Na and Cu contents in
mature sporophyte were much higher than in gameto-
phyte, and theNA of 12 nutrients inmaturesporophyte
followed the order: K ( 17 .7% ) > P ( 15 .1% ) >
Cu ( 13 .3% ) > N ( 11 .6% ) > Na ( 10 .5% ) >
Mn ( 7 .8% ) > Zn ( 7 . 5% ) > C ( 6 . 9% ) > Mg
(6 . 8% ) > Ca ( 5. 4% ) > Fe ( 1 . 3% ) > Al
(1. 2% ) , which indicates that there was a high de-
mand for these 5 nutrients in the developing sporophyte
of D. diversiformis . Mn was also essential to thedevel-
oping sporophyte of D. diversiformis . However, Fe and
Al contents were much lower in sporophyte than in ga-
metophyte, suggesting there was a little demand for
them . Therefore, we assumed that if the needs for N,
P, K , Na, Cu andMn can not besatisfied dueto lack
of precipitation during the reproductive seasons or due
to failure of competition with gametophyte, then the
failure of sexual reproduction in D. diversiformis may
occur .
Acknowledgements : This study was supported by the Center of
Analysis and Test of Wuhan University for nutrients analysis and
the Nature Protection Bureau of Fanjing Mountain for field assis-
tance . We thank Songquan SONG, Institute of Botany, Chinese
Academy of Sciences, for reading an early draft and offered nu-
merous suggestions, and Gongxi CHEN for offering constructive
criticisms on the manuscript .
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