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Asian Herpetological Research 2012, 3(2): 163–172
DOI: 10.3724/SP.J.1245.2012.00163
1. Introduction
Amphibians, considered good indicators of ecosystem
health, are experiencing population declines and many are
approaching extinction globally and rapidly (Stuart et al.,
2004; Wake and Vredenburg, 2008). Threats contributing
to the decline include habitat loss, alien species, over-
exploitation, global climate change, environmental
chemicals and emerging infectious diseases (Collins
and Storfer, 2003; Xie et al., 2007). At least 43.2%
of amphibian species have experienced some form of
population decrease between 1980 and 2004, compared
to the 27.2% which have remained stable and only 0.5%
which have experienced an increase (Stuart et al., 2004).
Endemic Amphibians and Their Distribution in China
Junhua HU, Cheng LI, Feng XIE and Jianping JIANG
*
Chengdu Institute of Biology, Chinese Academy of Sciences, Chengdu 610041, Sichuan, China
*
Corresponding author: Prof. Jianping JIANG, from Chengdu Institute
of Biology, Chinese Academy of Sciences, Sichuan, China, with
his research focusing on the systematics, evolutionary biology and
conservation genetics of amphibians.
E-mails: jiangjp@cib.ac.cn
Received: 21 December 2011 Accepeted: 10 May 2012
Endemic species are those entirely restricted to a specied
region and, on average, usually possess a narrower
geographic range than for a given taxonomic group
(Laffan and Crisp, 2003). It has been suggested that such
species have a high risk of extinction by chance alone
(Gaston, 1994; Myers et al., 2000), as they are conned
to limited geographic ranges and their localized habitats
are easy perturbed or destroyed, or they tend to be scarce
within their ranges (Brooks et al., 2002; Zhang, 2011).
Thus, they become one of the most effective surrogates
for identifying conservation priorities or hotspots, and
should be urgently targeted for conservation, management
and exploration of the underlying mechanisms of
biogeographic patterns (Myers et al., 2000; Sodhi et al.,
2008; Hu et al., 2011).
Endemic species (hereafter endemics) are not randomly
distributed within their distribution ranges, but are often
agglomerated in specific regions or habitats (Kluge and
Kessler, 2006). Due to their spatial distributions being
Keywords amphibian diversity, conservation, distribution pattern, elevational gradient, endemic species
Abstract Amphibians are good indicators of ecosystem health. Updating the information of endemic species in time
and understanding their spatial distributions are necessary for the development of integrative conservation strategies
and the elucidation of geographic patterns of amphibians. We analyzed the diversity and distributions of the endemic
amphibians in China based on a database of specimen records, recently published literature, and eld surveys. Two
hundred and sixty two endemic species of amphibians, belonging to 59 genera in 12 families and two orders, are
recognized. They account for 67% of the total number of amphibian species in China, with the family Megophryidae
possessing the largest number of endemics. There are also 17 genera endemic to China. Across provinces, the species
richness of endemics shows ve levels with a hierarchical diversication. Sichuan has the largest number of endemics,
while Heilongjiang and Jilin have no endemic species. Alternatively, whether on the level of endemic genus or species,
the highest diversity occurs in the Western Mountains and Plateau Subregion. The distribution ranges of most endemic
species cover ≤ 4 provinces or zoogeographic subregions. Additionally, the species richness of endemics along an
elevational gradient shows a bell-shaped pattern with the peak around 800 m. Most endemic species are distributed
in areas of low to mid elevation (c. 700–1 500 m), while 36 species are distributed up to 3 000 m. Endemic species
richness decreases with increasing of elevation range. Species at mid elevations display large range sizes, while species
at lower and higher elevations exhibit smaller ranges. Our results are to be beneficial for further exploration of the
underlying mechanisms of distributional patterns of amphibians in China. This study highlights a need to promote
conservation programs for Chinese endemic amphibians due to their narrow distribution ranges and potential threats.
Asian Herpetological Research164
Vol. 3
important to our understanding and conservation of
biodiversity, it has become a critical issue to analyze
species richness and endemism using observational and
specimen data (Myers et al., 2000; Crisp et al., 2001;
Hopper and Gioia, 2004; Hu et al., 2011). Additionally,
the formation of endemics is complicated and closely
related to geology, climate, and the process of bio-
evolution. Therefore, studies on endemics and their
distribution patterns are of great importance to learn about
the traits, components, origin and evolution of the local
fauna, and are also critical in studies of biodiversity and
conservation (Jiang, 1997).
China is considered as one of the important
biodiversity centers in the world (Myers et al., 2000;
Zhang, 2011). With its large area, diversied topography,
and varied climates and habitats, China has a high
degree of species diversity and endemism (Xie et al.,
2007; Zhang, 2011). However, percentages of vertebrate
endemics in China are uneven across taxonomic groups
(Fei et al., 2010a; Zhang, 2011). In contrast with the
endemism levels of about 8% in birds (Lei et al., 2003)
and about 18% in mammals (Smith and Xie, 2008),
current knowledge points to higher level of amphibian
endemism with 63% (Jiang et al., 2010). Moreover, most
endemic amphibians are rare and endangered or with
great significance in phylogeny and systematics (Xie et
al., 2007; Fei et al., 2010b; Hu et al., 2011). Chen et al.
(2008) also indicate that amphibians seem to be a better
indicator of zoogeographic division than either mammals
or birds.
Taxonomic revision and recognition of new species
of amphibians has occurred substantially in China (e. g.,
Fei et al., 2010b; Nishikawa et al., 2011a, b; Yang et al.,
2011). Presently, 389 amphibian species are recognized
in this country, belonging to 78 genera in 13 families and
three orders (Jiang et al., 2010; Hu and Jiang, unpublished
data). The importance of endemic amphibians to planning
the conservation of biodiversity has been emphasized
(Xie et al., 2007; Fei et al., 2010a). Although previous
studies related to endemic amphibians in China are
scatteredly mentioned in literature (e. g., Xie et al., 2007;
Fei et al., 2010a; Zhang, 2011), endemism has not been
spatially explored in any detail at the national scale
(Jiang et al., 2010), except for regional amphibians (Fu
et al., 2006). Moreover, endemic species numbers have
been underestimated in these studies due to the omission
of some species and recognition of new species after
being published and the inconsistent opinions in validity
of some species among herpetologists. To understand
the diversity and spatial distributions of the endemic
amphibians in China, we aim to: 1) revise the checklist
of endemic amphibians, 2) reveal their geographic
distributions at a regional scale (i. e., administrative
provinces and zoogeographic realm), and 3) document
the elevational patterns of species richness. Our study
will not only help to understand the general mechanisms
for distributions of Chinese endemic amphibians, but also
guide conservation planning for these endemics.
2. Methods
2.1 Database Although it is difcult to estimate the exact
geographic range of a single species, we can conrm the
provinces or zoogeographic realms where amphibians
species occur. We generated a spatial database of the
distribution of endemic amphibians in China as the basis
for this study. Data on geographic distributions were
obtained from the following sources: 1) specimen records
from museums, mainly the Herpetological Museum of
Chengdu Institute of Biology (CIB), Chinese Academy
of Sciences (CAS); 2) amphibian and/or herpetological
fauna (Fei et al., 2006, 2009a, b; Yang and Rao, 2008; Shi
et al., 2011); 3) herpetological monographs and current
literature (Zhao and Adler, 1993; Nishikawa et al., 2009,
2011a, b; Fei et al., 2010a; Yang et al., 2011); and 4)
the results of our field surveys during the 1970s–2000s
by the members of the Evolution and Biodiversity
Conservation Research Group of CIB, CAS. Species
taxonomy mainly followed Fei et al. (2006, 2009a, b,
2010a, b) and partially followed Frost (2011). The recent
changes in the classification of amphibians based on
phylogenetic relationships were fully considered and
some modications were made (Jiang et al., 2010).
We recognized endemics referenced in Fei et al. (2006,
2009a, b, 2010a) and Frost (2011), which represents
those only known to occur in China until now. The
data of elevational distributions (minimal and maximal
elevation of occurrences) for each species were compiled
through an exhaustive search of the primary literature
and museum collections, and complemented with field
records.
2.2 Statistical analysis The map of China (1:4 000 000)
was digitized to produce a geographic units (provinces)
map using ArcGIS 9.2 (ESRI, Redland, USA). It was
then overlaid with distribution maps to determine the
distributions of endemics in each province. Because most
records of endemics were documented for provinces but
not for municipality cities and special administrative
regions, we incorporated Tianjin and Beijing into Hubei
Province, Shanghai into Jiangsu Province, Chongqing
Junhua HU et al.
Endemic Amphibians and Their Distribution in China
No. 2 165
into Sichuan Province, and Hongkong and Macao into
Guangdong Province when analyzing. China covers
parts of the Oriental and Palaearctic zoogeographic
realms, including seven regions: Northeastern Region,
Northern China Region, Inner Mongolia-Xinjiang
Region, Qinhai-Xizang Region, Southwestern Region,
Central China Region and Southern China Region.
As each of these regions consists of 2–5 subregions
(Zhang, 2011), we compiled the species richness for each
subregion. Additionally, we considered a species to have
a narrow distribution if its range covers ≤ 4 provinces or
subregions, while a wide distribution was considered to
be one that covers ≥ 4 provinces or subregions (Jiang et
al., 2010).
To examine the relationship between species richness
of amphibians and elevation, we divided the elevation into
100 m bands and calculated the number of species in each
band. The mean and the difference between the minimum
and maximum elevations of occurrence reported for each
species were used to represent its elevational midpoint
and breadth. To overcome statistical non-independence
of the spatial data, we used the ‘midpoint method’ as a
measure of the central tendency (Hu et al., 2011). We
explored the relationships between the species richness
of endemics and their altitudinal ranges using the simple
ordinary least squares (OLS) model by Origin 7.5
(OriginLab Corporation, Northampton). P ≤ 0.05 was
considered statistically significant. Furthermore, for all
endemic amphibians, two-dimensional histograms were
used to depict the continuous variation of elevation range
profiles for range maxima (upper limit) and minima
(lower limit) over 500 m intervals, and for the midpoints
and range sizes over 500 m intervals along the altitudinal
gradient.
3. Results
3.1 Endemism of amphibians Two hundred and sixty
two endemics of amphibians are recognized, belonging
to 59 genera in 12 families and two orders (Appendix 1).
They account for 67% of the total species of amphibians
in China. Across families, Megophryidae possesses the
largest number of endemics (65 species), accounting
for 25% of all endemic amphibians. It is followed by
Ranidae and Rhacophoridae with 64 and 33 endemics,
respectively. Considering the endemics of zoogeographic
realms, there are more oriental species than palearctics.
244 species are oriental, occupying 93% of all endemics,
while only eight species are palearctic, and 10 species are
distributed in both the Oriental Realm and the Palaearctic
Realm.
There are 17 genera endemic to China, including
Batrachuperus Boulenger, 1878; Dianrana Fei, Ye
and Jiang, 2010; Feirana Dubois, 1992; Glandirana
Fei, Ye and Huang, 1990; Hypselotriton Wolterstorff,
1934; Liua Zhao and Hu, 1983; Liuhurana Fei, Ye,
Jiang, Dubois and Ohler, 2010; Liurana Dubois, 1986;
Oreolalax Myers and Leviton, 1962; Pachyhynobius
Fei, Qu and Wu, 1983; Pachytriton Boulenger, 1878;
Parapelophryne Fei, Ye and Jiang, 2003; Protohynobius
Fei and Ye, 2000; Pseudohynobius Fei and Ye, 1983;
Pseudorana Fei, Ye and Huang, 1990; Rugosa Fei, Ye
and Huang, 1990; and Yerana Jiang, Chen and Wang,
2006 (Appendix 1). For these genera, 17 species are
belonging to the genus Oreolalax, with 6 species in both
Pachytriton and Hypselotriton, 5 in both Pseudohynobius
and Batrachuperus, 4 in Liurana, 3 in Feirana, and 2
in both Liua and Pseudorana. The other genera are all
monospecic genus.
3.2 Distribution in provinces and zoogeographic
regions The abundance of endemics in provinces can be
divided into ve levels with a hierarchical diversication,
that is, Level I with 44–87 species, including Sichuan,
Yunnan, Hunan, Guizhou and Guangxi having a higher
diversity of 87, 63, 48, 45 and 44 species, respectively;
Level II with 21–30 species, including Guangdong,
Hubei, Fujian, Anhui, Zhejiang, Gansu, Xizang, and
Taiwan; Level III with 10–19 species, including Hainan,
Jiangxi, Henan, Shannxi and Jiangsu; Level IV with
1–6 species, including Qinghai, Hebei, Shandong, et al.;
Level V with no endemics, including Heilongjiang and
Jilin. Among the provinces in Level IV, Xinjiang, Inner
Mongolia and Liaoning all have only one species of
endemic amphibians (Figure 1).
At the level of genus, both Sichuan and Yunnan have
the highest diversity with 31 genera. Moreover, the
hierarchical order from highest to lowest for genus is
Sichuan/Yunnan > Guizhou > Hunan > Hubei > Guangxi
> Zhejiang… and that for family is Yunnan > Sichuan
> Guangxi > Guizhou > Hubei > Anhui > Guangdong...
(Figures 1). There is no endemic amphibian family in
China.
Alternatively, among zoogeographic subregions,
whether at the level of genus or species, the Western
Mountains and Plateau Subregion, Southwest China
Mountains Subregion, and Eastern Plain and Upland
Subregion have higher diversity than any of the others.
In the sequence, there are 77, 74 and 56 species within
these subregions, respectively. The Southern Yunnan
Subregion follows with 31 species. Other subregions have
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1–25 species, while the Daxing’an Mountains Subregion,
Changbai Mountains Subregion and South China Sea
Archipelago Subregion have no known endemics. At
the level of family with endemics, the Southern Yunnan
Subregion has the highest diversity, followed by the
Southwest China Mountains Subregion, and Western
Mountains and Plateau Subregion (Figure 2).
There are 232 (89% of the endemic amphibians in
China) and 258 (98%) endemics whose distribution
ranges cover ≤ 4 provinces and subregions, respectively.
Within them, 155 and 211 species only occur in a
single province or subregion. For example, Feirana
kangxianensis, Rana kunyuensis, Scutiger ningshanensis,
Pseudepidalea taxkorensis and Hynobius chinensis are
only found in Gansu, Shandong, Shannxi, Xinjiang
and Hubei, respectively. Xenophrys huangshanensis
and Pachytriton feii are both only distributed in Anhui;
Echinotriton chinhaiensis, H. yiwuensis and H. amjiensis
are all only in Zhejiang; Amolops daiyunensis, Odorrana
huanggangensis, Glandirana minima and Hypselotriton
fudingensis only in Fujian; and more other species, such
as O. kuangwuensis, X. wawuensis, and Batrachuperus
londongensis only in Sichuan, and H. wolterstorffi,
Oreolalax jingdongensis, and Feihyla fuhua only in
Yunnan. Only six species (2%) have the ranges covering
≥10 provinces. Across provinces, 34 endemics restrict
their distributions to Yunnan, followed by Sichuan with 27
species. For the endemic genera, only Batrachuperus and
Feirana are distributed in the Palaearctic Realm partially
and all the other genera are endemic to the Oriental
Realm. Specifically, Protohynobius, Parapelophryne,
Glandirana and Liurana are restricted to Sichuan,
Hainan, Fujian, and Xizang, respectively (Figure 1).
3.3 Elevational patterns of species richness and range
size Endemic amphibians of China are distributed over
a large spectrum of elevation from 0–4 500 m. The
relationship between the species richness and elevational
gradient shows a bell curve pattern with a normal
distribution (Figure 3). Endemics increase in number
steeply when elevation increases from 100 m to 800 m
and then reach the peak. Whereafter, endemics decrease
with the elevation increasing from 800 m to 2 800 m, and
only about 30 species occur in an elevation band ranging
Figure 1 Distribution patterns of endemic species of amphibians in
different provinces of China. The numbers in parentheses represent,
respectively, the number of genus and family that the endemic
species belong to.
Figure 2 The abundance of endemic species of amphibians in
different zoo-geographic subregions of China. The numbers in
parentheses represent, respectively, the number of genus and family
that the endemic species belong to.
I: Northeastern China Region (A:
Daxing’an Mountains Subregion; B: Changbai Mountains Subregion;
C: Songliao Plain Subregion); II: Northern China Region (A: Yellow
River-Huai River Plain Subregion; B: Loess Plateau Subregion); III:
Inner Mongolia-Xinjiang Region (A: Eastern Grass Land Subregion; B:
Western Arid Subregion; C: Tianshan Mountains Subregion); IV: Qinhai-
Xizang Region (A: Qiangtang Plateau Subregion; B: Qinghai-Southern
Xizang Subregion); V: Southwestern China Region (A: Southwest China
Mountains Subregion; B: Himalaya Mountains Subregion); VI: Central
China Region (A: Eastern Plain and Upland Subregion; B: Western
Mountains and Plateau Subregion); VII: Southern China Region (A:
Coastal Fujian, Guangxi and Guangdong Subregion; B: Southern Yunnan
Subregion; C: Hainan Island Subregion; D: Taiwan Island Subregion; E:
South China Sea Archipelago Subregion). The A–E after I–VII represent
subregions, following Zhang (2011).
Figure 3 The species richness of endemic amphibians along the
elevational gradients in China.
Junhua HU et al.
Endemic Amphibians and Their Distribution in China
No. 2 167
from 2 800 m to 3 300 m. Endemics further decrease and
nally only one species can be found at the elevation up
to 4 500 m (Figure 3).
Seventy eight endemics (about 30% of the Chinese
endemic amphibians) are distributed over large elevational
ranges (≥ 1 000 m). Among them, Andrias davidianus, A.
mantzorum, Batrachuperus tibetanus, Bufo bankorensis
have elevation ranges over 2 500 m. 36 endemics (14%)
are distributed at and above 3 000 m. A. davidianus,
B. karlschmidti, B. tibetanus, B. yenyuanensis, B.
tibetanus, Nanorana pleskei, N. ventripunctata, Scutiger
glandulatus, and Scutiger mammatus have the highest
elevation distribution over 4 000 m. On the other hand,
Echinotriton chinhaiensis, Hyla zhaopingensis, H.
yiwuensis, Nidirana hainanensis, Rana kunyuensis, O.
macrotympana, Pelophylax plancyi, Sylvirana hekouensis
and X. brachykolos are typical species with low elevation
(≤ 400 m). The distribution patterns also show that most
of the endemics occur in areas of low to middle elevations
ranging from 700 m to 1 500 m.
Correlation between the richness of endemics and the
elevation range is well explained by a simple ordinary
least squares (OLS) model (R
= 0.725, P = 3.495E-8;
Figure 4). Additionally, the continuous variation shown
by elevation range profiles can be made discrete for
range maxima and minima for all endemics ending at
4 500 m (Figure 5). Most endemics (79%) are depicted as
reaching their range maxima at low to middle elevation
(≤ 2 500 m), and very few surpass the elevation of 4 000
m. Ninety two endemics (35%) exhibit the combination
of range maxima and minima. Relatively large numbers
of endemics exclusive to highlands (the cell farthest to
the right) and diminishing numbers of species that extend
from highlands to lower elevation zone are clear traces
of a distinctive highland amphibians. A great proportion
of endemics (85%) have their lower limits below 2 000
m,
while only five species are with their lower limits
above
3 000 m.
The relationship between the range size and midpoint
of the elevation range reveals a triangular distribution for
Chinese endemic amphibians (Figure 6). The endemics
at mid elevations display the complete range sizes, while
those at lower or higher elevations possess only small
range sizes.
4. Discussion
4.1 Endemic amphibians and their distributions in
China Owing to the many high mountains and deep
river valleys along with the diversified landforms
and climates across a vast area, China has varied and
complicated ecological environments, which has enabled
amphibians to adapt to the diverse habitats and thus form
the unique distribution patterns (Fei et al. 2006, 2009a,
b). The identification of new species has increased the
number of endemic amphibians in China over the past
several decades (e. g., Zhao and Adler, 1993; Fei et
Figure 4 Correlation scatter-plots for species richness with the
elevational range for endemic amphibian species in China.
Figure 5 Correlation scatter-plots for species richness with the
elevational range for endemic amphibian species in China.
Figure 6 Two-dimensional histograms of elevational range limits
for endemic amphibian species in China. The color spectrum
represents the elevational gradients.
Asian Herpetological Research168
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al., 2008; Nishikawa et al., 2011b; Yang et al., 2011).
For example, Xie et al. (2007) recorded 215 species of
endemic amphibians, Fei et al. (2010a), 233 endemics
and presently we have recognized 262 endemics in
China. In this study, we present the first systematic
analysis of endemic amphibians in China, which would
give us a better understanding of the endemism and the
distributions of amphibians in China.
Previous biogeographic studies on amphibians in
China mainly focused on threatened species, diversity
across all species, or elevational patterns of regional
frogs (Fu et al., 2006; Xie et al., 2007; Jiang et al.,
2010). It is suggested that the level of available survey
efforts, the consequent number of records available for
assessment, and the identification of new species can
strongly influence biogeographic assessments (Slatyer
et al., 2007). Molecular analyses are increasingly used
in systematics and some species complexes or cryptic
species are recognized (e. g., Nishikawa et al., 2011b;
Yang et al., 2011). Concurrently, substantial eld efforts
for amphibians have been launched in recent years in
China. Such efforts have dramatically increased the
records available for analyzing the distributions of
amphibians and providing information of unsurveyed
or poorly surveyed areas. Jiang et al. (2010) indicated
that the greatest species richness occurred in the family
Ranidae, followed by Megophryidae. However, in this
study, Megophryidae is found with the largest number of
endemics, and Ranidae is the second. This discrepancy
may be due to the different distribution characteristics
between these two families: the proportion of species with
narrow distributions in the Megophryidae is higher than
that in the Ranidae. Among provinces, Sichuan is shown
to have the largest number of threatened amphibians
(Xie et al., 2007) and endemic birds (Lei et al., 2003),
while Heilongjiang and Jilin are located in the region of
low species diversity (Xie et al., 2007). This pattern is
coincident with that of endemic amphibians uncovered
here.
We can define the endemics which are limited to
small, narrow regions or in discontinuous regions as
species with regional or discontinued distributions (Lei
et al., 2003). For example, O. jingdongensis is only
distributed in Jingdong, Shuangbai, and Xinping counties
of Yunnan, even if there are wide similar habitats around
their known distribution sites (Fei et al., 2009a). The
difference between the ‘actual distribution areas’ and
‘potential distribution areas’ may inevitably occur (Hu
and Jiang, 2010). Except for some determined new
differentiation patterns, the explanation for the locally
distributed endemics is mostly human activities or
that the species are at a natural decline state (Hu and
Jiang, 2010; Zhang, 2011). Additionally, the isolated
discontinuous distribution is also a normal phenomenon
of local distribution. For example, H. orientalis is widely
distributed from eastern coastal to the central mainland of
China, but its congener, H. chenggongensis, can only
be
encountered in Chenggong County, Yunnan (Fei et al.,
2006; 2010a). Another example is the genus Feirana. F.
quadranus is widely distributed in the Wuling Moutains,
Daba Mountains, Shennongjia, Longmen Mountains,
and the central and western Qinling Mountains, while F.
taihangnica in the central and eastern Qinling Mountains,
Zhongtiao Mountains, and southern Taihang Mountains
(Wang et al., 2009; Yang et al., 2011; Hu and Jiang,
unpublished data). The central Qinling Mountains are
suggested as the sympatric areas between these two
species of Feirana (Wang et al., 2009). However, F.
kangxianensis, which is closely related to F. taihangnica,
is currently known to only occur in Kangxian County,
Gansu, and is discontinuously distributed in those areas
of F. taihangnica (Yang et al., 2011). The mainland,
Taiwan Island and Hainan Island of China have similar or
related species, which are also discontinuously distributed
with a representative example of the genus Hynobius.
H. arisanensis, H. formosanus, H. fuca, H. glacialis,
and H. sonani are endemic to Taiwan Island, while H.
amjiensis, H. guabangshanensis, H. maoershanensis,
and H. yiwuensis occur in Anji County of Zhejiang,
Qiyang County of Hunan, Longshen and Xing’an
counties of Guangxi, and Zhenhai, Yiwu, Wenling,
Jiangshan, Xiaoshan and Zhoushan counties of Zhejiang,
respectively (Shen et al., 2004; Fei et al., 2006, 2010a;
Zhou et al., 2006; Lai and Lue, 2008). These distribution
patterns may relate to the appearance of several ice ages
in the Quaternary or the vicariance events (Fei et al.,
2006; Zeisset and Beebee, 2008).
4.2 Elevational patterns of species richness and range
size The understanding of the distribution patterns of
species richness along elevational gradients is crucial to
developing a general theory on species diversity (Rowe,
2009). Our results show that the endemic amphibians in
China are not evenly distributed along the elevational
gradients, with the richest species being at elevations
between 800–1 200 m. Looking at a wide geographic
range for frog taxa, Hu et al. (2011) suggested that mid-
elevations (around 1 500 m) in Asia are richer in endemics
of spiny frogs than the lower and higher elevations. In
this study, the substantial decrease of endemic amphibians
towards high elevations (> 3 000 m) further supports this
Junhua HU et al.
Endemic Amphibians and Their Distribution in China
No. 2 169
pattern. Similar patterns are also documented in the frogs
of the Hengduan Mountains (Fu et al., 2006). It can be
interpreted as a reflection of the complex topographic
configuration of China, where many montane habitats
possess varied and complicated ecological environments.
Accumulations of endemics are likely to be found in the
highly diversified habitats. This fact indicates that the
amphibian community composition pattern is strongly
inuenced by the basins with stepped landforms (Fei et al.,
2009b; Zhang, 2011).
Distribution ranges of species result from complex
interactions among many factors, including physiological
traits, history of speciation and dispersal, and constraints
from continental shape (Webb and Gaston, 2003). The
fact that the richness of endemic amphibians in China
decreases with increasing elevational range indicates
that most endemics possess narrow elevational ranges.
In particular, many endemics exhibit the combination of
lower and upper limits. Here, a triangular pattern is found
for the relationship between range sizes and midpoints
of elevation, and species at intermediate elevations
have the broadest amplitudes. The results complement
other evidences showing a need for further testing the
generality of the Rapoport’s altitudinal effect (Patterson et
al., 1998; Hu et al., 2011).
5. Conclusions
Endemism is inherently scale dependent and sensitive
to the delineation of boundaries (Lomolino et al.,
2006). Therefore, we should delimit the distributions
of endemic taxa ideally using natural and geographic
boundaries rather than administrative boundaries.
Moreover, two types of errors (i. e., the overweighting
of the widespread species regarded in the literature but
with few observational records; the erroneous records
for species with narrow recorded ranges which create
comparatively high endemism scores in wrong places),
should be emphasized when the distributions of endemics
are being interpreted (Slatyer et al., 2007). Despite
these issues, exploring the patterns of endemism and the
distributions of endemics is important for understanding
regional characters, compositions and evolutionary
process of fauna, and also is of particular interest in
the development of integrative conservation strategies
(Jablonski, 1986; Grau et al., 2007). Amphibians
with restricted ranges should be urgently protected
(Sodhi et al., 2008). Therefore, China which is one of
the countries with richest endemic species needs to
pay immediate attention to conservation (Xie et al.,
2007). For example, it is urgently necessary to launch
conservation plans urgently for F. kangxianensis, given
its restricted distribution range and great existence risk
under the pressure of intense human activities (Yang
et al., 2011; Hu and Jiang, unpublished data). Besides
the endemic amphibian species, there are 17 endemic
genera in China with some monospecific genera. They
are especially important for the evolutionary process and
need conservation, because the extinction of one species
means that all of the genus will disappear. Unfortunately,
China’s natural habitats, particularly forests, have suffered
severe degradation because of increasingly intensive
human activities (Xie et al., 2007; Jiang et al., 2010).
Thus, we still use administrative boundaries to delimit
endemic amphibians and refer to endemic taxa as those
only occurring within China’s borders. Alternatively,
we can use alternative methodologies for generating
distribution maps and range sizes for the endemics, such
as using predictive spatial modeling which would reduce
the problem of an undersampled species range (Hu et al.,
2010; Hu and Jiang, 2011). To be brief, this study can
promote our understanding of the underlying mechanisms
of distribution patterns of amphibians in China, and
highlights a need to launch conservation programs
for Chinese endemic amphibians due to their narrow
distribution ranges and the potential threats.
Acknowledgments We are grateful to Prof. Liang FEI,
Prof. Changyuan YE, Prof. Xiaomao ZENG, Dr. Yuchi
ZHENG, Dr. Bin WANG, Dr. Jiatang LI, and Dr. Li
DING for providing support and help in collecting the
information of endemic amphibians in China. We thank
Jing CHEN, Raul E. DIAZ, and Chunlan ZHANG for
help with earlier versions of the manuscript. We also
thank the two anonymous referees and the editors for their
invaluable suggestions. This study was supported by the
Knowledge Innovation Program of the Chinese Academy
of Sciences (Y1B302100) to Junhua HU; the National
Natural Science Foundation of China (NSFC, 31071906),
and the Main Direction Program of the Knowledge
Innovation Project of CAS (Grant No. KSCX2-EW-J-22)
to Jianping JIANG.
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