全 文 ： 菌物学报
jwxt@im.ac.cn 15 March 2013, 32(2): 216-225
Http://journals.im.ac.cn Mycosystema ISSN1672-6472 CN11-5180/Q © 2013 IMCAS, all rights reserved.






Supported by Flora of the Cryptogamics of China (NSFC grant 31093440), National Natural Science Foundation of China (No. 30770012),
Beijing Natural Science Foundation (No. 5123044) and Beijing Academy of Science and Technology Menya Project [2009].
*Corresponding author. E-mail: hmanrong@yahoo.com.cn, guosy@im.ac.cn
Received: 14-03-2012, accepted: 23-05-2012

Altitudinal patterns of the lichen genus Cladonia (Lichenized
Ascomycota) in China
GUO Shou-Yu1 DENG Hong1 BI Hai-Yan2 XIA Xiao-Fei2 HUANG Man-Rong2*
1State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing 100101, China
2Beijing Museum of Natural History, Beijing 100050, China



Abstract: Altitudinal patterns of the lichen genus Cladonia and 18 species in the genus were investigated based on
herbarium collections. Species in the genus are inclined to grow at high elevation. Normal and lognormal probability patterns
are ubiquitous for vertical distribution of these species, and vertical patterns are somewhat elevation-dependent. The
altitudinal ranges of the species were found to decrease with increase of altitude. These evidences suggest an immediate
threat from global warming to some species growing at higher elevation in the genus. Ecological evidence also confirms C.
bacillaris and C. macilenta are conspecific.
Key words: probability distribution models, unimodal patterns, Rapoport’s rule, fruticose lichen

中国石蕊属地衣的垂直分布规律
郭守玉 1 邓红 1 毕海燕 2 夏晓飞 2 黄满荣 2*
1中国科学院微生物研究所真菌学国家重点实验室 北京 100101
2北京自然博物馆 北京 100050
摘 要：基于标本馆馆藏标本的信息，研究了中国石蕊属 Cladonia 地衣及其 18 个种的垂直分布规律，并确认了本属
倾向于分布在高海拔地区的特点。物种随海拔梯度的变化在统计学上呈正态分布或偏正态分布是非常普遍的现象，
但是它们的分布规律与其所处的海拔位置相关：物种分布范围随着海拔的增加而缩小。因此，全球变暖会对高海拔
地区的物种构成更直接的威胁。粉杆石蕊 C. bacillaris 和瘦柄红石蕊 C. macilenta 具有相同的海拔分布规律和分布范
围，从生态学的角度支持了它们为同种的观点。
关键词：概率分布模型，单峰模式，拉波波特法则，枝状地衣


DOI:10.13346/j.mycosystema.2013.02.013
GUO Shou-Yu et al. / Altitudinal patterns of the lichen genus Cladonia (Lichenized Ascomycota) in China
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INTRODUCTION
Altitude is a key factor that influences
biodiversity (Sergio & Pedrini 2007). Today, within
the global warming context, the upward shifts of the
altitudinal zones of organisms, especially those
living in high elevations, have induced deep worries
in the conservation biology (Pounds et al. 2005).
Lichens, together with other cryptogamic plants,
have not been in the attention of mainstream of
conservation biology for a long time (Hunter &
Webb 2002). Therefore, it is not surprising that there
are no documented shifts of life zones for such
organisms at all. However, some studies on the
lichens of Livingston Island, South Shetland Islands,
Antarctica showed that lichens colonized soon after
the glacier retreat (Sancho & Pintado 2004; Sancho
et al. 2007). These results can be explained either by
an enlargement or shift of distribution areas of
lichens. Whatever may it be, the quick responses of
slow-growing lichens to global warming are
amazing. Altitude has effects on distribution of
organisms which is believed to mimic latitude
(Stevens 1992), thus, the investigation to altitudinal
patterns of lichens is also urgent. Only based on such
works, can the responses of lichens to global
warming be understood and can effective
conservation plans be made. Huang (2010) found
that some species of a common fruticose lichen
genus, Stereocaulon Hoffm. (1796), which is
inclined to grow on high elevation, statistically have
altitudinal patterns in the form of normal or
lognormal possibility distributions. This is also true
for the 11 species of Peltigera Willd. (1787) in
China (Liu et al. 2011). These results showed an
innate character of the distribution of organisms, and
may be referred by conservation biology to design a
conservation- and cost-effective plan to conserve the
biodiversity on our planet.
Cladonia P. Browne (1756) is the largest
fruticose lichen genus with more than 500 species
distributed world around (Kirk et al. 2008). Most
species of this genus are common in montane areas
and are also inclined to grow at high elevation,
which implies its sensitivity to global warming, and
thus urges us to perform an altitudinal analysis here.
1 MATERIALS AND METHODS
Instead of plots designation and samples
investigation, the source of elevation data in this
study was herbarium collections which have been
lodged in Herbarium Mycologicum Academiae
Sinicae-Lichenes (HMS-L) and were examined by
the first author. The merit and statistical reliability of
such data have been shown by other authors
(Hamilton 1975; Alcántara et al. 2002; Baniya et al.
2010; Huang 2010; Liu et al. 2011). To ensure the
accuracy of the analysis, the species with more than
5 specimens were considered in analysis of
altitudinal distribution pattern of the genus, and the
species with more than 50 specimens were analyzed
for the species altitudinal pattern.
The methods in Huang (2010) were followed
through. All elevation data were analyzed with both
their original forms and logarithmic forms to base
Euler’s number (e). The independent sample T-tests
were performed to test the equality of variances and
means of elevation data of the species using the
statistical software SPSS 16.0 (SPSS Incorporated,
Chicago, Illinois). The nonparametric tests with a
one-sample Kolmogorov-Smirnov test were
employed to determine the probability distribution
model describing the altitudinal pattern of each
taxon. All above-mentioned procedures were applied
to original and log-transformed data sets respectively.
Once a detailed distribution model was
established for a taxon, we calculated the altitudinal
range into which about 68% individuals were
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expected to fall according to the following steps: 1)
If the specified model was a normal probability
distribution, the expected elevational range was
between μ-σ and μ+σ, where μ and σ stand for the
mean and variance, respectively, and the average
elevation of this taxon was exactly the same to μ, the
arithmetic mean; 2) If the specified model was a
lognormal distribution, the expected range was
between exp(μ-σ) and exp(μ+σ), and the average
elevation was derived from calculating exp(μ)
instead of the arithmetic one. When the average
elevations of the taxa and their expected altitudinal
ranges were determined, we carried out the bivariate
correlation tests to examine if they were correlated.
We also grouped these species based on such
average elevation data using K-means cluster
process. The differences between these groups in
variances and means were tested using a One-way
ANOVA test.
2 RESULTS
2.1 Summary of the data set
A total of 2,813 specimens belonging to 115
species of Cladonia with unambiguous elevation
data on the tags were obtained, and 88%
(2,469/2,813) of them belonging to 104 species were
collected from the regions of altitude above 1,000m.
However, only 72 species each had no less than 5
specimens were adopted in the analysis of altitudinal
pattern of the genus. For 20 species each had no less
than 50 specimens, 18 of them were used to analyze
altitudinal pattern of species.
2.2 Independent sample T-tests for variances and
means of the 20 species
For original elevation data and logarithmically
transformed data, we got 190 species pairs for 20
Table 1 One-sample Kolmogorov-Smirnov tests for the elevation data of Cladonia and its eighteen species
Taxa N Probability distributions P Mean Std. deviation Altitude range (m)
C. alinii 50 Lognormal 0.159 7.650c 0.270c 1603.59–2751.77
C. amaurocraea 135 Lognormal 0.140 7.538c 0.498c 1141.39–3090.23
C. bacillaris 94 Lognormal 0.227 7.297c 0.707c 727.78–2992.91
C. chlorophaea 53 Normal 0.753 2061.32 973.31 1088.01–3034.63
C. coccifera 85 Normal 0.089 2727.99 961.49 1766.5–3689.48
C. coniocraea 75 Lognormal 0.374 7.421c 0.514c 999.25–2793.36
C. furcata 86 Lognormal 0.223 7.493c 0.480c 1110.98–2901.55
C. gracilisa 93 Normal 0.094 1402.90 443.04 959.86–1845.94
C. macilenta 64 Lognormal 0.101 7.157c 0.868c 538.61–3056.42
C. macropteraa 61 Normal 0.091 2801.18 696.49 2104.69–3497.67
C. ochrochlora 108 Normal 0.280 2315.09 842.43 1472.66–3157.52
C. pleurota 99 Uniform 0.109 2376.55 1273.83 -
C. pocilluma 71 Normal 0.277 2533.87 960.53 1573.34–3494.40
C. pyxidata 110 Normal 0.498 2603.54 831.50 1772.04–3435.04
C. ramulosa 100 Normal 0.326 1536.55 816.28 720.27–2352.83
C. scabriusculaa 118 - 0.042 1883.05 694.83 -
C. stellaris 57 Lognormal 0.090 7.594c 0.484c 1224.15–3222.78
C. yunnana 51 Normal 0.530 3370.39 523.72 2846.67–3894.11
Cladoniab 72 Lognormal 0.569 7.611c 0.310c 1481.78–2754.52
Note: aBased on the data set after the exclusion of redundant data; bBased on average elevation of the seventy-two species with no less than fifty
specimens; cCalculated from the logarithmically transformed data.
GUO Shou-Yu et al. / Altitudinal patterns of the lichen genus Cladonia (Lichenized Ascomycota) in China
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species in each data set. Among them, equivalence
of variances were supported for 71 species pairs for
original elevation data and 60 for logarithmically
transformed data, equivalence of means were
supported for 47 species pairs for original elevation
data and 44 for transformed data. While equivalence
of both means and variances were supported for 20
species pairs for original data and 21 for transformed
data, with 5 were strongly supported (F<1.000,
P>0.300) for original and transformed data
respectively. These results suggest that, while a
small fraction of these species occupy similar
elevations, most of them differ to each other
significantly in altitudinal distributions.
2.3 One-sample Kolmogorov-Smirnov tests for
the genus and its 18 species
When all data were included in the
Kolmogorov-Smirnov tests, we detected the
lognormal distribution for the genus Cladonia in
China (P=0.569, Table 1, Fig. 1), and suitable
probability distribution models for 14 species: the
normal distribution for 6 species, the lognormal
distribution for 7 species, and the uniform
distribution for 1 species (Table 1, Fig. 1).


Fig. 1 Histograms of Cladonia and its seventeen species with their altitudinal pattern curves. Only the genus and 17 species which were
detected with statistical distribution models were shown.
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Table 2 K-means clusters of the sixteen species with normal or lognormal distributed patternsa
Species Distribution patterns Group Distance to centre (m)
C. coccifera Normal I 2.647
C. macroptera Normal I 75.837
C. pyxidata Normal I 121.803
C. pocillum Normal I 191.473
C. ochrochlora Normal I 410.253
C. yunnana Normal I 645.047
C. ramulosa Normal II 182.528
C. gracilis Normal II 316.178
C. chlorophaea Normal II 342.242
C. coniocraea Lognormal II 48.375
C. furcata Lognormal II 76.352
C. amaurocraea Lognormal II 158.992
C. bacillaris Lognormal II 243.213
C. stellaris Lognormal II 267.164
C. alinii Lognormal II 381.567
C. macilenta Lognormal II 436.022
Note: aTwo groups were obtained: group I’s center is 2,725.34m a.s.l., while group II’s center is 1,719.08m a.s.l. One-way ANOVA test of these two
groups suggested that F=38.196, P<0.001.
When redundant data were ruled out, a further
analysis of the other 4 species showed that 3 species
[C. gracilis (L.) Willd., C. macroptera Räsänen and
C. pocillum (Ach.) O.J. Rich.] have the normal
probability distribution models (Table 1, Fig. 1; see
below in Discussion part 3.2).
2.4 K-means clusters of the species
Cluster tests for the 16 species with normal or
lognormal distribution models (Table 1, Fig. 1)
resulted in 2 groups (Table 2, Fig. 2). Group I had 6
species that were normally distributed, with cluster
center at 2,730m, and group II had the other 10
species, the 7 lognormally distributed species and
the other 3 normally distributed ones, with cluster
center at 1,720m. These two groups differed
significantly in variances and means (F=38.196,
P=0.000) in one-way ANOVA test.

Fig. 2 Results of K-means clusters of the sixteen Cladonia
species. The cluster center of group I is 2,730m, that of group II is
1,720m. (▲) Normally distributed species; (■) Lognormally
distributed species.
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2.5 The bivariate correlation test of the average
elevations and altitudinal ranges
The bivariate correlation test for the 13 species
with normal or lognormal probability distributions
(when all data were included, see Table 1) show that
the altitudinal ranges and average elevations were
significantly and negatively correlated (r=-0.660,
P=0.014; Fig. 3-A). However, when incorporating
C. gracilis, C. macroptera and C. pocillum with
normal probability distribution models (detected
when redundant data were ruled out, Table 1) into
the model, this negative correlation became weaker
and less significant (r=-0.370, P=0.158; Fig. 3-B).

Fig. 3 Correlation of the averaged elevations and altitudinal
ranges of the species with normal or lognormal patterns. A: The
thirteen species with normal or lognormal patterns detected when
all data were included (Pearson correlation coefficient r=-0.660,
two tailed significance value P=0.014); B: Sixteen species with
normal or lognormal patterns when the redundant data were ruled
out (r=-0.370, P=0.158).
3 DISCUSSION
3.1 The distribution of Cladonia in China
Phytogeographic studies in lichenology are
dependent both on sound taxonomic records and
detailed distribution maps (Goward & Ahti 1997). In
China, the number of species of the Cladonia varies
largely in different regions. Eighty-nine species,
constituting 72% of the total number of the species
in China, occur in south-western China, an area
marked by widely stretching mountain ranges, 62
(50%) occur in mountainous northeastern China,
especially Changbai Mountain and Da Hinggan
Mountains, 55 (45%) in Qingling Mountains in
central China, 39 (32%) in eastern China, and 37
(30%) in tropical Guangxi (Guo 1999), respectively.
Floristic and chemical diversity of the Cladonia in
China are greater at relatively higher and forested
areas than lower areas (Guo 2000). Our results also
suggested that overwhelming majority of the
investigated species (104/115) can be found in the
regions of altitude above 1,000m. This fact
confirmed previous reports.
Huang (2010) spotted a normal distribution
model to another fruticose lichen genus
Stereocaulon, and confirmed the high-elevated
distributions of that genus. In this study, we also
found a unimodal altitudinal pattern of Cladonia
(Fig. 1) and recognized lognormal instead of normal
distribution for it in Kolmogorov-Smirnov test
(P=0.569, Table 1). According to the obtained
empirical function, the average elevation of this
genus is 2,020m and 68% of species are distributed
between 1,480m and 2,750m (Table 1). This fact
suggests that this genus is also a high-elevated genus
and we must keep a close watch on its dynamics in
the context of global warming.
But little is known about the life history and
ecology of most species in Cladonia, especially for
the species with much branched thalli and lacking
reproduction structures (apothecia and/or pycnidia).
Some of them were catalogued as endangered, for
example, the amphipacific endemic species C.
pseudoevansii Asahina (Wei et al. 1986). In China, it
only appears in south slope of Changbai Mountain in
Jilin Province between 1,500 and 2,200m a.s.l. This
causes concerns regarding its recolonization
potential, since small fragments may not disperse
either far or fast. To maintain the habitat and current
environment in the narrow region may be a good
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choice for protecting C. pseudoevansii.
The altitudinal pattern analysis of species may
yield valuable suggestions for applied management
of the endangered species. Further studies of boreal
and temperate forest terrestrial Cladonia species
biology and ecology may offer useful information
for management of the terrestrial lichen
communities including some key species.
3.2 The altitudinal patterns of the 18 species of
Cladonia
Different altitudinal patterns have been
described by various authors. While some authors
detected that species diversity declines with
increasing elevation (MacArthur 1972; Rohde
1992), some others concluded that the highest
species diversity occurs at the mid-elevations (Bruun
et al. 2006; Pinokiyo et al. 2008). This fact suggests
that different functional and taxonomic groups have
different altitudinal patterns (Grytnes et al. 2006). In
our studies, unimodal patterns were detected for 13
species of Cladonia when all data were included,
among which 6 species were distributed following
normal probability distribution functions, and the
other 7 following lognormal ones (Table 1, Fig. 1).
The only exceptional species, C. pleurota (Flörke)
Schaer., was found to be uniformly distributed
according to elevation gradient (Table 1, Fig. 1),
which has never been spotted in other taxa before.
This result makes a further confirmation that
different taxa follow different altitudinal patterns,
though most species seem to have a unimodal
pattern.
The other 6 species each had no less than 50
specimens in the original data set have not been
detected any probability functions to describe them.
We revisited the data set carefully and found that a
large number of specimens of each species with the
same elevation were collected in the same localities
by the same collectors. We assumed that these data
induced noises and had an important effect on the
analyses, being the same effect of repeating data
from a single plot for several times in sample
investigations. Therefore, for each species, we left
one of such data to represent the corresponding
elevation and ruled out the others, then applied the
re-assembled data set to Kolmogorov-Smirnov test
again. After such artificial selection of data, only 4
species remained more than 50 individual elevation
data, viz. C. gracilis, C. macroptera, C. pocillum
and C. scabriuscula (Delise) Leight. Results show
that the first 3 species were subject to normal
probability distribution according to altitude (with
P=0.094, 0.091, 0.277 respectively, Table 1), while
the last species, C. scabriuscula, was not to any
probability distributions (P=0.042 in the normal
distribution test, Table 1). But such processes of data
exclusion are subjective somewhat and may also
cause deviations, and the extent of its influence
remains to be estimated.
Whatever may it be, these results combined
with that in Huang (2010) suggest that, though some
taxa (such as the case of C. pleurota in this study)
are exceptional, normal and lognormal probability
distribution are ubiquitous altitudinal patterns for
organisms in nature. In fact, we suppose that Grytnes
et al. (2006; Fig. 3-B therein) has detected a normal
distribution for the altitudinal pattern of vascular
plants in the Jondalen municipality in western
Norway and Baniya et al. (2010; Fig. 3 therein) for
lichens in Nepal though they have not been strictly
tested.
Huang (2010) hypothesized that the altitudinal
patterns of a certain taxa are elevation-dependent.
He found that those species of Stereocaulon growing
at low elevations were normally distributed, those
growing at middle elevation were lognormally
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distributed, while those growing at high elevation
were distributed irregularly. However, Liu et al.
(2011) found the normal altitudinal distribution
pattern for Peltigera species growing at higher
elevations, and the lognormally distribution pattern
for those species growing at lower elevations. We
examined the relationship between altitudinal
distribution pattern and average elevation in
Cladonia. The cluster analysis based on the average
elevation of the 16 species with normal or lognormal
altitudinal distribution patterns obtained 2 groups
which differed significantly in variances and means
(F=38.196, P=0.000, Table 2). Group I consists of 6
normally distributed species exclusively, while all 7
lognormally distributed species fall into group II
together with the other 3 normally distributed
species, viz. C. chlorophaea (Flörke ex Sommerf.)
Spreng., C. ramulosa (With.) J.R. Laundon and C.
gracilis (Table 2). Accordingly, these two groups are
corresponding to these two probability distribution
models on the whole. Group I, with cluster center of
2,730m, stands for the normally distributed species
at higher elevations, while groups II, with cluster
center of 1,720m, for the lognormally distributed
species at relatively lower elevations (Fig. 3).
Therefore, the altitudinal patterns of Cladonia
species, with a way being contrast to Stereocaulon
but similar to Peltigera, also seem to be
elevation-dependent. This fact suggests that these
elevation-dependent altitudinal patterns are common
to a certain extent; at least it is true for these two
typical fruticose lichen genera.
3.3 Altitudinal range of the species according to
the altitudinal gradient
It has been long recognized that species
richness decreases with increasing latitude, and
Stevens (1989) presented a second latitudinal
correlations which he called Rapoport’s rule,
describing a phenomenon that latitudinal ranges of
taxa increase with increasing latitude. Later, as
altitudinal effects to the distribution of organisms are
thought to mimic latitudinal effects, Stevens (1992)
extended Rapoport’s rule to altitude, and
demonstrated it with many examples.
However, what generality of Rapoport’s rule is
equivocal (Rohde 1996; Gaston et al. 1998). Rohde
(1992) found that marine teleost fishes have the
greatest latitudinal ranges at low latitude. This rule
was not suitable for either tree species or liverworts
in Nepal (Bhattarai & Vetaas 2006; Grau et al.
2007), and did not supported by computer
simulations (Stauffer & Rohde 2006).
We examined Rapoport’s rule with Cladonia
species here. At first, considering that the process of
ruling out redundant data for C. gracilis, C.
macroptera, C. pocillum and C. scabriuscula may
cause unexpected deviations, we carried out the test
for the other 13 species. According to the result, the
altitudinal ranges decrease with increasing altitude
(r=-0.660, P=0.014, Fig. 3-A). After adding data of
C. gracilis, C. macroptera and C. pocillum, a
decreasing trend of altitudinal ranges was still
detected though it was not so significant (r=-0.370,
P=0.158, Fig. 3-B). This obvious difference of these
coefficients and significance values may be
attributed to the artificial exclusion of elevation data
of the later 3 species. Anyway, decreasing trends of
altitudinal ranges were detected in both cases, and
thus the extension of Rapoport’s rule to altitude is
not supported here.
The concerns about the shifts of distributional
ranges of organisms (Pounds et al. 2005) with the
global warming would be intensified if the declining
trends of altitudinal ranges with increasing altitude
are proved to be common for endangered species in
high elevations, because the up-shifts of life zones
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will shrink their small distributional and make a
further challenge for the efforts of biodiversity
conservation.
3.4 The classification of C. bacillaris and C.
macilenta
For classification, a highly integrated science,
new evidences are always welcome, even if they
seem slightly significant. For the last two decades,
we’ve seen the overwhelmingly rise of molecular
data which have changed the taxonomy and
systematics profoundly. However, taxonomists and
systematists have never given up their effort to seek
other sources of data, such as ecological evidences
(Bondartseva & Komarov 2011).
In Cladonia, C. bacillaris (Ach.) Genth and C.
macilenta Hoffm. occur widespread in the whole
world. At present, many authors regarded them as
the same species, and accepted C. macilenta as the
correct name (Ahti 2000). Two main chemical races
(chemotype I: PD-, K-, KC+ yellow to orange, C-,
barbatic acid with or without squamatic acid, mainly
on soil; chemotype II: PD+ orange, K+ yellow, KC-,
C-, thamnolic acid with or without barbatic acid,
mainly on old wood and bark of tree), which were
often regarded as distinct species (C. macilenta and
C. bacillaris) before based on the presence or
absence of thamnolic acid and the different habitats,
may have subtle morphological and ecological
differences, at least in some regions. In our study,
these two species have the same altitudinal pattern
(lognormal probability distribution, Table 1), and the
equality of variances and means of the elevation data
sets were strongly supported in independent samples
t-tests (F=1.398, P=0.246 for original data; F=0.053,
P=0.265 for logarithmically transformed data).
Therefore, it is obvious that these two species
occupy the same altitudinal range. This evidence
provides another support to the conspecific status of
the two species. However, it is in need of further
taxonomical study worldwide with data from various
sources to establish their relationship.
4 CONCLUSION
The fruticose lichen genus Cladonia is inclined
to grow at high-elevated montane areas, while the
species richness of the genus changes according to
altitudinal gradients statistically following a
lognormal probability distribution and reaches its
peak at middle altitude. Most species investigated
here are normally or lognormally distributed,
depending on their averaged elevation, which
suggests that those unimodal patterns may be
ubiquitous in organism distributions. Rapoport’s rule
is not recognized here; instead, the higher elevation
a species of Cladonia grows at, the smaller
altitudinal range it occupies. Thus, the high elevated
life zones of this genus may very sensitive to global
warming and call for close monitoring to their
dynamics. Our analysis also supports C. bacillaris
and C. macilenta to be conspecific in an ecological
consideration.

Acknowledgement: We are grateful to Lin-De Liu in
Beijing Museum of Natural History for English
corrections. We also want to express our sincere thanks to
Dr. Hua-Jie Liu (Hebei University) for the constructive
suggestions, and to Dr. Lawrence A. Glacy (Brigham
Young University, Utah, USA) for the English corrections.
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