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The geographical distribution of grey wolves (Canis lupus) in China: a systematic review

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The grey wolf (Canis lupus) is one of the most widely distributed terrestrial mammals, and its distribution and ecology in Europe and North America are largely well described. However, the distribution of grey wolf in southern China is still highly controversial. Several well-known western literatures stated that there are no grey wolves in southern China, while the presence of grey wolf across China has been indicated in A Guide to the Mammals of China, published by Princeton University Press. It is essential to solve this discrepancy since dogs may have originated from grey wolfs in southern China. Therefore, we systematically investigated Chinese literatures about wild animal surveys and identified more than 100 articles and books that included information of the distribution of grey wolves in China. We also surveyed the collections of three Chinese natural museums and found 26 grey wolf skins specimens collected across China. Moreover, we investigated the fossil records of wolf in China and identified 25 archaeological sites with wolf remains including south China. In conclusion, with the comprehensive summary of Chinese literatures, museum specimens and fossil records, we demonstrate that grey wolves does distribute across all parts of the Chinese mainland, including the most southern parts of China.
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ZOOLOGICAL RESEARCH
Science Press Zoological Research 37(6): 315-326, 2016 315
The geographical distribution of grey wolves (Canis
lupus) in China: a systematic review
Lu WANG1,#, Ya-Ping MA1,2,#, Qi-Jun ZHOU2, Ya-Ping ZHANG1,2, Peter SAVOLAINEN3, Guo-Dong WANG2,*
1 State Key Laboratory for Conservation and Utilization of Bio-resources in Yunnan, Key Laboratory for Animal Genetic Diversity and
Evolution of High Education in Yunnan Province, Yunnan University, Kunming 650091, China
2 State Key Laboratory of Genetic Resources and Evolution, Yunnan Key Laboratory of Molecular Biology of Domestic Animals, Kunming
Institute of Zoology, Chinese Academy of Sciences, Kunming 650223, China
3 Science for Life Laboratory, Department of Gene Technology, KTH-Royal Institute of Technology, Solna 17165, Sweden
ABSTRACT
The grey wolf (Canis lupus) is one of the most widely
distributed terrestrial mammals, and its distribution
and ecology in Europe and North America are
largely well described. However, the distribution of
grey wolves in southern China is still highly
controversial. Several well-known western literatures
stated that there were no grey wolves in southern
China, while the presence of grey wolves across
China has been indicated in A Guide to the
Mammals of China, published by Princeton
University Press. It is essential to solve this
discrepancy since dogs may have originated from
grey wolves in southern China. Therefore, we
systematically investigated Chinese literatures about
wild animal surveys and identified more than 100
articles and books that included information of the
distribution of grey wolves in China. We also
surveyed the collections of three Chinese natural
museums and found 26 grey wolf skins specimens
collected across China. Moreover, we investigated
the fossil records in China and identified 25
archaeological sites with wolf remains including
south China. In conclusion, with the comprehensive
summary of Chinese literatures, museum specimens
and fossil records, we demonstrate that grey wolves
do distribute across all parts of the Chinese
mainland, including the most southern parts.
Keywords: China; Grey wolf; Distribution; Conservation
INTRODUCTION
The grey wolf, Canis lupus, is one of the most widely distributed
terrestrial mammals (Young & Goldman, 1944). Grey wolves
live in a wide variety of habitats, including the dry Arabian
desert, the xeric Mediterranean shrublands, the coniferous
forests of Siberia, and the frozen tundra on Ellesmere island
(Mech, 1981). Despite extirpation from many parts of their
previous range over the last few hundred years, by persecution
from humans and habitat fragmentation (Hunter & Barrett, 2011;
Young & Goldman, 1944), wolves still retain most of their
original distributions.
The distribution and ecology of grey wolves are largely well
described in Europe and North America. However, in more
peripheral and remote parts of their distributions, detailed
information is often lacking. In the western literature, the wolf
has generally been reported to be distributed throughout the
northern hemisphere, from N15° latitude in North America and
N12° latitude in India to beyond the Arctic Circle, but has been
considered to be absent from Africa and the southern East Asia
(Mech, 1981). However, recent articles reported that the
Egyptian jackal (Canis aureus lupaster, Hemprich and
Ehrenberg 1833) was not a subspecies of the golden jackal
(Canis aureus, Linneaus 1758) and should be reclassified as
the African wolf, Canis lupus lupaster (Gaubert et al., 2012;
Koepfli et al., 2015; Rueness et al., 2001). 1
Similarly, the literature about wolves in China is limited
outside China. This has led to misconceptions in the
western literature about the distributions of wolves in
Received: 01 November 2016; Accepted: 10 November 2016
Foundation items: This study was supported by grants from the
National Natural Science Foundation of China (91531303), the 973
program (2013CB835200 and 2013CB835202), the Breakthrough
Project of Strategic Priority Program of the Chinese Academy of
Sciences (XDB13000000), and grants from the Carl Trygger
Foundation and the Agria and Swedish Kennel Club research
foundation. GD Wang is supported by the Youth Innovation Promotion
Association, Chinese Academy of Sciences
#Authors contributed equally to this work
*Corresponding author, E-mail: wanggd@mail.kiz.ac.cn
DOI:10.13918/j.issn.2095-8137.2016.6.315
www.zoores.ac.cn 316
China. Four studies, all conducted by western researchers,
stated that wolf has never been presented in large parts of
China (Callaway, 2013; Larson & Fuller, 2014; Nowak, 2003;
Sokolov & Rossolimo, 1985 ).
However, as will be shown in this study the grey wolf has a
historical and current range across nearly the entire country of
China. There are more than 100 Chinese articles and books
involving investigations of wolves in China since the 1950s
(Table 1), showing the distributions in detail. Most of these
articles are species investigations at a provincial or local level,
however, there is no comprehensive description of the current
distribution of wolves across China. Therefore, we here
summarized the Chinese literature concerning past and
present distributions of wolves in China, in order to
synthesize data from this rich source of regional
investigations into a comprehensive map of wolf distribution in
China, and to make this significant information available to
an international audience.
Table 1 Literature list of distributions of wolves in China
Province Investigation year Location Reference
Heilongjiang 2008-2009 Eastern forests of Wandashan Mountains Shen et al., 2011
1994-2001 Sanjiang National Reserve Zhang et al., 2001
1997-1999 Tangwanghe river forest distict He et al., 2003
1993-1995 Grand Khingan Zhang et al., 1998a
1988-1989, 1993-1995 Northern Grand Khingan Li et al., 1996
1984, 1987-1990, 1992 Xingkai Lake Nature Reserve Li et al., 1993
1971-1980, 1981-1990 Heilongjiang province Zhang et al., 1998b
N/A Heilongjiang province Zhang & Yu, 2005
N/A Western Helongjiang province Gao et al., 1999
Jilin 1992-1993 Yanbian Qiu et al., 1995
1992-1993 Hunjiang Li et al., 1994
Liaoning 1999-2002 Nuluerhusan National Reserve Zhou et al., 2007
1996-2000 Benxi Zhao et al., 2004a
1996-2000 37 counties in Liaoning province Zhao et al., 2004b
1996-1999 Fushun Zhao et al., 2001
N/A Yiwulv Mountain National Nature Reserve Liu et al., 2008
N/A Liaoyang Wang et al., 2004
Inner Mongolia 1985-1986 Jiufeng Shan Liu & Liu, 1999
N/A Chaihe Xiao et al., 2013
N/A Hulunbair & Hinggan Gao et al., 1999
Beijing 1982-1983 Changping & Miyun Zhang, 1984
N/A Beijing Wu et al., 2006
Tianjin N/A Tianjin Wu et al., 2006
Shanxi 2010-2011 Pangquangou National Nature Reserve Wang & Zhao, 2011
1996-1997 Luyashan Nature Reserve Qiu et al., 1998
N/A Northeastern Loess Plateau Chen, 2000
Hebei 1993-2001 Chengde Hou et al., 2004
N/A Hebei province Wu et al., 2006
N/A Saihanba Hou et al., 1994
Gansu 2007-2009 Sunan and Subei prairie Zhao et al., 2011
N/A Gannan plateau Chen & Li, 1994
N/A Longnan mountain Chen et al., 1994
N/A Tianshui Hu et al., 1993
N/A Minqin desert Chen, 1992
N/A Anxi Chen & Luo, 1991
Zoological Research 37(6): 315-326, 2016 317
Continued
Province Investigation year Location Reference
Xinjiang 1994-1996 Kanas National Nature Reserve Abdukadi et al., 1999
1987-1988 Wuqia, Taxkorgan, Yecheng, Qiemo, Yutian Feng, 1990
1965, 1980, 1983, 1985 Zhungeer & Altai Zhang & Hu, 1988
1979 Xinjiang Gao, 1997b
1958-1961 Desert plains area in Xinjiang Zhang, 1963
N/A West Tianshan National Nature Reserve Liu et al., 2007a
Ningxia 2010-2011 Luoshan National Nature Reserve Qin & Chang, 2012
Shaanxi 2006 Huanglongshan Nature Reserve Li & Liu, 2009
2006 Micangshan Nature Reserve Wen et al., 2008
1997-2000 Changqing National Nature Reserve He, 2001
1999 Zhashui Hu et al., 2003
1996 Zhouzhi National Nature Reserve Li & He, 1997
1963-1966 Ankang Wu & Li, 1982
1959 Daba mountain Wang et al., 1981
N/A Shaanxi province Li et al., 2006
Qinghai 2001-2002 Qilian mountain Xia et al., 2003
N/A Beichuan River Nature Reserve Zhang & Pu, 2012
N/A Qinghai lake area Kong et al., 2011
Tibet 2001-2002 Upper Zayu river basin Wu, 2006
1987-1988 Ngari & Naqu Feng, 1990
Sichuan 2006 Kasha Lake Nature Reserve Liu et al., 2013
1997, 2006 Ruoergai Wetland National Nature Reserve Liu et al., 2009
2005-2006 Maozhai Nature Reserve Liu et al., 2007b
2003-2005 Haizishan Nature Reserve Liu et al., 2007c
2004 Heizhugou Nature Reserve Liu et al., 2005a
2002-2003 Jiuzhaigou National Nature Reserve Liu et al., 2005b
2002-2003 Dafengding Nature Reserve Liu et al., 2004
2002-2003 Yele Nature Reserve Zhang & Hu, 2004
2001-2002 Huanglong Nature Reserve Zhu et al., 2010
2002 Xuebaoding Nature Reserve Sun et al., 2006
2001 Pingwu He et al., 2004
1998 Big-small Langou Nature Reserve Lu & Hu, 2003
1996 Huanglongsi Nature Reserve Hu et al., 2001
N/A Ganzi and Liangshan Zhang et al., 2009
N/A Ruoergai Wetland National Nature Reserve Hao et al., 2008
N/A Wolong Nature Reserve Yu et al., 1983
Yunnan 2010-2011 Lanping Yunling Provincial Nature Reserve Cui et al., 2014
2010-2011 Weixi Zha et al., 2014
N/A Yunnan province Yang et al., 1999
Guizhou 2005-2006 Leigong Mountain National Nature Reserve Chen et al., 2008
N/A Guizhou province Luo & Li, 2001
N/A Weining Huang, 1989
www.zoores.ac.cn 318
Continued
Province Investigation year Location Reference
Chongqing 2006-2008 Jinfo Mountain Natural Reserve Zong et al., 2010
1995 Jinfo Mountain Natural Reserve Peng et al., 1996
N/A Chongqing Han & Hu, 2002
Henan 1997 Xin'an, Yuzhou, Jiyuan, Luoning, Jiaozuo, Zhenping Gan & Fan, 2004
Hubei 2004 Yerengu Nature Reserve Wang et al., 2007
2004 Wudaoxia Nature Reserve Wu et al., 2005
2001 Qizimei Mountain Nature Reserve Liu et al., 2002
N/A Duheyuan Provincal Nature Reserve Li et al., 2008
Hunan 1980-1981 Ziyunshan Fu, 1987
Jiangxi 2004-2007 Taohong Ridge Sika Deer Nature Reserve Wu et al., 2012
1984-1986 Poyang lake area Fu & Ding, 1991
N/A Jiangxi province Tu et al., 2014
N/A Lushan Nature Reserve Li et al., 2007
Shandong 1984-1987 Jiaodong peninsula Sun, 1988
1982-1986 Qingzhou Cong, 1988
1961-1966, 1973-1984 Jiaodong and Luzhongnan area Lu, 1984
N/A Laoshan Tian et al., 2000
Anhui 1959-1964 Anhui province Wang et al., 1966
N/A Anhui province Wu et al., 2002
N/A Huangshan Xu, 1997
Jiangsu N/A Jiangsu province Wang & Zhao, 2008
Zhejiang 2005-2008 Hangzhou Ding et al., 2008
1958-1960, 1962-1964, 1979-1981 Zhejiang province Zhuge, 1982
N/A Jinhua Zhu & Yu, 1996
N/A Yongkang Bao & Hu, 1987
Fujian N/A Fujian province Chen et al., 2009
N/A Fujian province Zhou, 1997
N/A Fujian province Zhan, 1995
Guangxi 1997-2000 Shiwan Mountain Xia et al., 2002
1958 Southwestern Guangxi Wang et al., 1962
Guangdong 2000 Nanling National Nature Reserve Fellowes et al., 2003
LITERATURE SUMMARIZATION
It is controversial to describe the distribution of grey wolves in
western literatures. Two articles reported that wolves were
previously present all across China, but is now extinct from
southern China (Ginsberg & Macdonald, 1990; Lau et al., 2010).
In four well-known studies, researchers claimed that wolves have
never existed in sourthern China (Callaway, 2013; Larson &
Fuller, 2014; Nowak, 2003; Sokolov & Rossolimo, 1985),
suggesting that sourthern China cannot be the harbor of dog
domication. Thus, southern China is usually treated outside the
range of wolf distribution (IUCN; EOL). However, in 2008, Smith
and his colleagues described the distribution of wolf in China,
indicating that grey wolves were present all across the mainland
of China (Smith & Xie, 2008).
In the Chinese literature, wolves have been reported to
appear over all parts of continental China. The Fauna Sinica
(China): Mammalia Vol. 8 Carnivora page 46-49, reported in 1987:
“the wolf, which apart from Hainan Island, the various islands in
the South China Sea, and Taiwan, is spread over nearly all the
country“ and ”the wolf can be seen in all provinces. Based on
collected literature references and specimen samples, wolves
have been identified in Muleng, Baoqing, and Genhe of
Heilongjiang, in Baicheng, Kaitong, Dunhua, Jingyu, Huinan,
Hunchun, Jilin, Tumenling, and Fuyu of Jilin, in Fushun and Lvda
of Liaoning, in Shanhaiguan and Zhangjiakou of Hebei, in Beijing,
in Hohhot and Erlian of Inner Mongolia, in Hami, Bole, Turpan,
Zoological Research 37(6): 315-326, 2016 319
Yanqi, Korla, Aksu, Luntai, and Baicheng of Xinjiang, In Shanxi
province, in Yan’an of Shaanxi, in Mianchi and Luoning of Henan,
in Yichang of Hubei, in Nanjing and Qingjiang of Jiangsu, in
Fujian province, in Longzhou, Ningming, and Shangsi of Guangxi,
in Guangdong province, in Guizhou province, in Lushui and
Chengkou of Yunnan, in Yumen, Zhangye, and Linxia of Gansu,
in Menyuan, Qilian, Alaer, Golmud, and Delingha of Qinghai, in
Pali, Nylamu, Tingri, Shigatse, and Naqu of Tibet, and in Shiqu,
Ruoergai, Songpan, Leibo, Ebian, Kangding, Wanxian, Yibin, and
Mianyang of Sichuan” (Gao & Wang, 1987).
Furthermore, Wang (2003) described the subspecies/
subtypes of grey wolvesin China and reported that they were
distributed across all parts of continental China. Chinese wolves
were divided into five subspecies and forms: Canis lupus
desertorum Bogdanow, 1882 in Xinjiang, C. l. filchneri Matschie,
1907 in Qinghai, Gansu and Tibet, C. l. chanco Gray, 1863 in
Heilongjiang, Jilin, Liaoning, Inner Mongolia (eastern part),
Hebei, Beijing, Shandong, Henan and Shanxi, C. l. Nei-Mongol
form in Inner Mongolia (western and mid part) and C. l. South-
China form in Anhui, Jiangsu, Zhejiang, Jiangxi, Fujian,
Guangdong, Hunan, Guizhou, Yunnan, Hubei and Sichuan
In order to obtain an updated and comprehensive description
of the distribution of wolves in China, we investigated more than
100 articles containing information about the presence of wolf at
a regional level (see a full list of literature in Table 1). The most
recent evidence of wolf in each province (Figure 1) were
extracted from the following papers: Heilongjiang (Shen et al.,
2011), Jilin (Qiu et al., 1995), Liaoning (Zhou et al., 2007), Inner
Mongolia (Liu & Liu, 1999), Beijing (Zhang, 1984), Tianjin (Wu
et al., 2006), Shanxi (Wang & Zhao, 2011), Hebei (Hou et al.,
2004), Gansu (Zhao et al., 2011), Xinjiang (Abdukadir et al.,
1999), Ningxia (Qin & Chang, 2012), Shaanxi (Li & Liu, 2009),
Qinghai (Xia et al., 2003), Tibet (Wu, 2006), Sichuan (Liu et al.,
2013), Yunnan (Cui et al., 2014), Guizhou (Chen et al., 2008),
Chongqing (Han et al., 2010), Henan (Gan & Fan, 2004), Hubei
(Wang et al., 2007), Hunan (Fu, 1987), Jiangxi (Wu et al., 2012),
Shandong (Sun, 1988), Anhui (Wang et al., 1966), Jiangsu
(Wang & Zhao, 2008), Zhejiang (Ding et al., 2008), Fujian
(Chen et al., 2009), Guangxi (Xia et al., 2002), Guangdong
(Fellowes et al., 2003).
Figure 1 Distributions of wolves in China
The latest investigation year recorded in literature in 26 provinces (in red) and the latest publication year of literature in three provinces (in green) are
indicated within brackets.
www.zoores.ac.cn 320
In summary, these investigations showed that the wolf has
been recorded in every continental Chinese province between
1964 and the present, except in three provinces (Figure 1 in
green). Most notably, wolves were recorded in South China (in
Yunnan province) as late as 2011 and in the two southernmost
continental provinces (Guangdong and Guangxi) in the year of
2000. From these findings we concluded that wolves are still
present across all parts of continental China.
WOLF SKINS IN ZOOLOGICAL MUSEUMS
In addition to the literature investigation, we made a survey of
wolf skins in the archives of the National Zoological Museum of
China, Kunming Natural History Museum of Zoology, and
Shaanxi Institute of Zoology, and (Table 2, Figure 2, Figure 3).
We found 26 wolf skins sampled from 13 provinces across
China, e.g., two specimens sampled from two southern
Chinese provinces (Zhejiang and Fujian) in 1974, and one from
southern Yunnan in 1985.
WOLF FOSSIL RECORD
We investigated the literature about archaeological research in
China, to identify information about wolf fossils in archaeological
sites. We extracted information about the fossil record of the
grey wolf in China from three Chinese books (Lv, 2004; Yuan,
2015; Zhang et al., 2003). These books reported 25 archaeological
sites in 14 provinces across China with wolf fossils records
(Table 3), including the 12 000 years old remains from the
South Chinese province Jiangxi.
Table 2 Sources and geographical origins of wolf skin specimens
Museum ID Province Location Date
1 Heilongjiang Baoqing N/A
2 Heilongjiang Baoqing 1957.01.24
3 Inner Mongolia Xiguitu (Yakeshi) 1954.12.10
4 Jilin Baicheng 1957.02.11
5 Jilin Jingyu 1956.03.08
6 Jilin Kaitong 1956.06.13
7 Xinjiang Buerjin 1974
8 Xinjiang Bole 1972.05.18
9 Tibet N/A N/A
10 Tibet Changdu 1976.1
11 Tibet N/A N/A
12 Beijing Yanqing 1984.04.28
13 Sichuan Ruo’ergai 1961.07.03
14 Yunnan Lushui 1960
15 Fujian N/A 1974.05
The National Zoological Museum of China, Beijing
16 Zhejiang Lin’an 1974
17 Yunnan Kunming 1967
18 Yunnan Kunming 1957
19 Yunnan Zhaotong N/A
20 Yunnan Honghe 1985
21 Guizhou N/A N/A
22 Guizhou N/A N/A
Kunming Natural History Museum of Zoology, Kunming
23 Jiangxi Zoo 1990.06.08
24 Shaanxi Yan’an 1973
25 Shaanxi Xunyang 1965
Shaanxi Institute of Zoology, Northwest Institute of
Endangered Zoological Species, Xi’an
26 Shaanxi Pingli 1965
DISCUSSION
In this study, we showed that contrary to what is reported in
many references in the western literature, the grey wolf actually
is present across virtually all parts of the mainland China. This
correction is important in studies of wolf ecology and
conservation. It gives a correct picture of the worldwide
Zoological Research 37(6): 315-326, 2016 321
Figure 2 Source and geographical origin of museum wolf skin specimens
distributions of wolves, by filling in a large blank region on the
map. It is also important in studies of the history of domestic
dogs, since dogs probably trace a large proportion of their
genetic ancestry to wolves from the southern parts of East Asia
(Wang et al., 2016).
The wolf has endured massive decline in population size and
geographic range around the world during the previous two
centuries, because of human influence including habitat loss,
persecution, hunting (for obtaining, e.g., trophies, furs and
material for traditional medicine), and depletion of prey
(Beschta & Ripple, 2010; Callan et al., 2013; Levi & Wilmers,
2012; Ripple et al., 2014). Also in China, the distribution areas
of wolves have severely decreased due to human mediated
habitat loss and hunting (Gao, 1997a, 2006; Zhang, 1999).
Official investigations from the middle of the 20th century
reported that wolves were distributed in every province of China
except some islands, but gave no exact numbers. Today, large
populations remain only in the northwestern and northeastern
parts of the country, Inner Mongolia and Tibet, but even in these
regions, the numbers are relatively small, e.g., only 2 000
wolves in Inner Mongolia were reported in the 1990s (Gao,
1997a). We have here shown that wolves still seem to be
present across all parts of the Chinese mainland, including the
most southern provinces. Thus, even though habitat loss has
been severe in urban and agricultural regions, wolves seem to
have persisted in intervening regions.
The data about wolf distributions that we here present were
investigations on either provincial or local level, whereas, a
comprehensive ecological survey of the wolves in China. It is
therefore not clear how the wolf populations in the different
parts of China are interrelated. For example, it is not clear
whether wolves recorded in the southern provinces represent
permanent populations, or a steady stream of individuals
migrating from the northern provinces. However, it is notable
www.zoores.ac.cn 322
Figure 3 Three museum wolf skin specimens
Specimens originating from Yunnan Province (left, ID 18 in Table 2), Jiangxi Province (middle, ID 23 in Table 2) and Shaanxi Province (right, ID 24 in
Table 2).
Table 3 Fossil records of gray wolves
Province County Archaeological site Time Reference
Shanxi and Hebei Yanggao and Yangyuan Xujiayao About 100 000 years ago Zhang et al., 2003 p259
Shaanxi Pucheng Nanwan and Beiwan Epipleistocene p315
Henan Anyang Xiaonanhai 22 150-11 000 years ago p320
Heilongjiang Harbin Yanjiagang 22 370±300 years ago p357
Shanxi and Hebei Yanggao and Yangyuan Xujiayao 125 000-104 000 years ago Lv, 2004 p96
Hebei Yangyuan Banjing 108 000-74 000 years ago p100
Shanxi Yanggao Shenquansi 11 720±150 years ago p102
Liaoning Haicheng Xiaogushan Epipleistocene p207
Chongqing Fengjie Yufupu 7 560±110 years ago p355
Heilongjiang Mishan Xinkailiu 7 500-6 500 years ago Yuan, 2015 p114
Qiqihar Tengjiagang Bronze age p115
Hailin Xilinhe Bohai Kingdom (698-926 A.D.) p115
Jilin Nong’an Zuojiashan 6 800-4 800 years ago p115
Liaoning Dalian Guojiacun 5 780-4 300 years ago P118
Inner Mongolia Linxi Baiyingchanghan 8 000-5 000 years ago P120
Baotou Yanjialiang 1 275-1 372 years ago p127
Shaanxi Nanzheng Longgangsi 6 500-6 000 years ago p130
Tongchuan Beicun Shang Dynasty (1 600-1 046 B.C.) p133
Hebei Xushui Nanzhuangtou About 10 000 years ago p144
Beijing Fangshan Zhenjiangying and Tazhao Shang and Zhou Dynasties (1 600-256 B.C.) p145
Shandong Yanzhou Wangyin 6 500-5 500 years ago p147
Weifang Qianbuxia
Houli Culture (8 500-7 500 years ago)
and 5 500-5 000 years ago p147
Tibet Naqu Chaxiutang 9th-11th century A.D. p155
Hubei Zigui Liulinxi
Neolithic age, Erlihe Culture (21st-15th
century B.C.), and the Eastern Zhou
Dynasty (770-256 B.C.)
p158
Badong Lijiatuo Eastern Zhou Dynasty (770-256 B.C.) p164
Jiangxi Wannian Xianrendong About 12 000 years ago p166
Zoological Research 37(6): 315-326, 2016 323
that wolves have been recorded across virtually the entire
continental China, including southern Chinese province Yunnan
as late as in 2011 and provinces Guangdong and Guangzhou in
2000. These findings indicate a consistent presence of
permanent populations across southern China. Moreover, to
obtain a comprehensive picture of the status of the wolves in
China, it is necessary to carry out both ecological and genetic
studies, e.g., in concerning the genetic relationships either
among the wolf populations across China and between these
and worldwide wolf populations.
This study points out misconceptions in the western literature
about the distributions of wolves in China. The origin of this
problem is not clear, but it can be traced back as far as an
article in 1985 from which the factoid has, stepwise, been
passed on to other articles (Sokolov & Rossolimo, 1985). It is
probably because of the linguistic barrier to the Chinese
literature that this error has previously not been pointed out.
This case can be explained by inefficient research in peripheral
parts of the species distribution, in countries with limited
resources. Our study raises the question whether this kind of
misconceptions also exist in other species than just the grey wolf.
CONCLUSIONS
With a comprehensive summary of Chinese literature, specimens
and fossil records, we showed that wolves are present across
all parts of the Chinese mainland, including the southern parts.
Hereby we corrected an error in western literature, in which
most sources stated that wolves were not present in the
southern China, and some even claimed that wolves have
never been presented there, even in ancient times. There is no
comprehensive description of the current distributions of wolves
across China, and therefore this study serves both to give an
updated description of wolf distributions in China, and to make
this significant information available to an international audience.
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... Within the Holarctic gray wolf complex, the highest diversity is found in wolves from Europe, China, and Russia (Ersmark et al., 2016). Focusing on Asia, Wang et al. (2016) described five wolf taxa for China, but the supporting evidence is scarce: C. l. chanco, C. l. filchneri, and C. l. desertorum (in Table 1 listed according to their contemporary use), C. l. Nei-Mongol form in Inner Mongolia (western and mid part) and C. l. ...
... NCBI GenBank currently lists C. l. chanco as the Mongolian wolf (NCBI GenBank Taxonomy Canis lupus chanco, 2019) and separately C. l. laniger as the Tibetan wolf (NCBI GenBank Taxonomy Canis lupus laniger, 2019). Wang et al. (2016), in their review on wolves in China, used C. l. chanco according to past (now outdated) usage, i.e., using C. l. chanco for the wolf lineage in Mongolia and northern China. They described this subspecies in the Chinese provinces of Heilongjiang, Jilin, Liaoning, Inner Mongolia (eastern part), Hebei, Beijing, Shandong, Henan, and Shanxi, but these populations may belong to C. l. lupus. ...
... al. (2013a) provides full mtDNA of one animal (used inFigure 2) but no phylogenetic researchZhang et al., 2013a;Wang et al., 2016 ...
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... Gray Wolf range in China is underrepresented on the Range Map of IUCN Red List (seeWang et al., 2016 ). ...
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We investigated the distributional range and population status of Hog deer through two-year camera trapping in Southwestern Yunnan Province and found that Hog deer have been extirpated from China due to habitat loss and illegal hunting. We recommend establishing a protected area in Nanting river watershed, to restore habitat and to reintroduce individuals from range of countries as the conservation priorities of the Regionally Extinct Hog Deer in China.
... As a result of management following national and international conventions and in spite of control efforts, wolves are recovering in many regions (Bragina et al., 2015;Chapron et al., 2014;Mech, 2017;Nickel & Walther, 2019). Still, the area of suitable habitat, especially in the southern parts of Eurasia, continue to decline and human pressures continue to increase (Kaczensky et al., 2008;Olson & Fuller, 2017;Reading et al., 1998;Wang et al., 2016). ...
Article
Wolf predation on domestic animals is a main reason for human-wolf conflict throughout the global range of wolves (Canis lupus). We conducted research in Daursky State Nature Biosphere Reserve and Valley of Dzeren Nature Refuge to evaluate the extent of wolf-livestock conflict. We documented 64 cases of livestock predation by wolves between 2015 and 2019 and analyzed the patterns of conflicts and people’s attitude towards wolves. A total of 283 livestock were killed by wolves, with an annual mean of 55.4 (SD = ± 1.44) animals/yr and a mean frequency of attack of 12.8 ± 3.89 attacks/yr. Sheep were the main prey of wolves, comprising 77.4 % of the total number of livestock killed. Among cattle and horses, wolves preferred to kill juveniles rather than adults (χ2 = 140.2, df = 2, P < 0.001). We found no significant difference between type of livestock killed by wolves by season (χ2 = 5.53, df = 3, P = 0.4776). Predations only occurred in situations where there were no protective actions, like a shepherd or protective corral. The mean annual rate of livestock lost by predation was 0.281 % (SD = ± 0.007). Attitudes towards wolves were mostly neutral to negative. The higher the level of income estimated by respondents (y = -0.242x + 0.98; R² = 0.99; F = 203.3; P = 0.005), the less negative the attitude towards wolf. To effectively reduce depredation, we suggest improving management actions, especially increased surveillance. We also discuss other management measures to mitigate livestock depredation.
... Western literature has incorrectly stated that wolves are absent from today´s Southern East Asia, with some articles claiming that wolves were never present in this region, even in ancient time. However, this was recently corrected in a review of the distribution of the wolf in China [21], showing that the southernmost record of wolf in modern time is in Yunnan in South China, only 200km north of Thailand. This suggests a theoretical possibility that the domestic dog may have originated as far south as Thailand. ...
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The dog population of Southern East Asia is unique in harboring virtually the full range of the universal mtDNA gene pool, and consequently, it has the highest genetic diversity worldwide. Despite this, limited research has been performed on dog genetics within this region. Here we present the first comprehensive study of a sub-region within Southern East Asia, analyzing 528bp of mtDNA for 265 dogs from Thailand, in the context of dogs from across the Old World. We found that Thailand was the only region in the world that has the full range of the universal mtDNA gene pool, that is, all 10 sub-haplogroups. Consequently, the statistics for diversity are among the highest, especially in North Thailand, which had high values for haplotype diversity and the number of haplotypes, and the lowest proportion of individuals with a universal type-derived haplotype (UTd) among all regions. As previously observed, genetic diversity is distinctly lower outside Southern East Asia and it decreases following a cline to the lowest values in western Eurasia. Thus, the limited geographical region of Thailand harbors a distinctly higher genetic diversity than much larger regions in western Eurasia, for example, Southwest Asia and Europe which have only five and four of the 10 sub-haplogroups, respectively. Within Thailand, diversity statistics for all four sub-regions follow the general pattern of Southern East Asia, but North Thailand stands out with its high diversity compared to the other regions. These results show that a small part of Southern East Asia harbors the full range of the mtDNA gene pool, and they emphasize the exceptional genetic status of Southern East Asia. This indicates that today's dogs can trace a major part of their ancestry to Southern East Asia or closely situated regions, highlighting Thailand as a region of special interest. Considering the large genetic diversity found in Thailand and that many neighboring regions, e.g., Myanmar and Laos, have not been studied for dog genetics, it is possible that large parts of the dog gene pool remain undiscovered. It will be an important task for future studies to fill in these blanks on the phylogeographic map.
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Several viruses can infect wild carnivores but their impact on wildlife health is poorly understood. We investigated the presence, diversity and distribution of various DNA viruses in 303 wolves inhabiting a vast area of the Northwest Territories, Canada, over a period of 13 years. We found evidence for the presence of canine bufavirus (CBuV, 42.6%), canine parvovirus 2 (CPV-2, 34.0%), canine bocavirus 2 (CBoV-2, 5.0%), cachavirus (CachaV-1, 2.6%), canine adenovirus 1 (CAdV-1, 1%) and minute virus of canines (MVC, 0.3%). To our knowledge, this is the first detection of CBoV-2, MVC and CachV-1 in wild animals. We also demonstrate that CBuV and CachaV-1 were already circulating among wild animals at least 11 and 10 years, respectively, before their discoveries. Although CBuV prevalence was higher, CPV-2 was the most prevalent virus among juveniles, while CBuV infection was associated with poor nutrition conditions. Even if its prevalence was low, CachaV-1 had the highest multiple infection rate (87.5%). CadV-1 and MVC sequences were highly identical to reference strains, but we observed a high diversity among the other viruses and detected three new variants. One CPV-2 variant and one CBuV variant were endemic since the beginning of the 2000s in the entire investigated region, whereas one CBuV variant and two CBoV-2 variants were found in a more restricted area over multiple years and CachaV-1 was found only in one region. Two CPV-2 variants and one CachaV-1 variant were observed only once, indicating sporadic introductions or limited circulation. Different patterns of endemicity might indicate that viruses were introduced in the wolf population at different timepoints and that mixing between wolf packs may not be constant. Different epidemiological behaviors depend on viral factors like infectivity, transmission routes, pathogenicity and tissue-tropism, and on host factors like proximity to densely populated areas, carnivory and pack density and mixing.
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The hog deer Axis porcinus formerly occurred in south-west China but has not been recorded there since 1965. To investigate the current status of the species in China, we conducted interviews, and transect and camera-trap surveys during October 2018–June 2020 to search for signs of hog deer across its historical range in the country. We interviewed 50 local inhabitants and surveyed 14 line transects in four counties of Lincang City, Yunnan Province. The camera traps were deployed in Nangunhe Nature Reserve (39.4 km of transects, 82 camera stations, 15,120 camera days) and Daxueshan Nature Reserve (41.1 km of transects, 68 camera stations, 13,554 camera days). We found no hog deer tracks and no hog deer were trapped by cameras. The floodplain grasslands preferred by hog deer along Nanting River have been transformed into agriculture plantations and human settlements. Our findings suggest that hog deer may have been extirpated from China, most likely as a result of habitat loss and overhunting. The conservation priorities for this species in China are the establishment of a protected area in the Nanting River watershed, restoration of habitat and reintroduction of individuals from range countries.
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Annapurna Conservation Area (ACA) is highly potential and domineering ecological habitats for wildlife and fauna. The region is home to elusive wild animals such as Himalayan wolf, Snow leopard and Tibetan fox. Himalayan Wolf (Canis lupus chanco) is the ancient linage of the dog-clade which is historically inhabiting in the Nepalese Himalaya, which has been extirpated from the most of its historic range since a couple of decades. Majority of the people in the region depends on livestock herding and tourism. Maximum Entropy (MAXENT) model was used to find the potential habitat of Himalayan wolf in Upper mustang, ACA. Presence-only data were collected throughout the feasible 2*2 Km2 grids. To get the better result 13 environmental variables were computed. Raster images were worked in ArcGIS and EDRAS Imagine and computed in MAXENT. The Naïve occupancy of Himalayan wolf was estimated 71.87%. Relative Availability Index and average entropy values were 2 sign samples/ km and 7.929 respectively predicting the whole study area potential wolf habitat. 1089 km2 area is estimated as potential Himalayan wolf habitat out of 2076.56 km2 studied. The study also shows high habitat overlapping between carnivores and livestock. MAXENT model, has shown high level of predictive performance with an ‘Area Under Curve’ between 0.876±0.983 and average Standard Deviation 0.044. It is understood that AUC value closer to 1, more suitable the model against computed predictive variables whereas lesser the value of Standard Deviation, less chance of overfitting among the presence data. The model has revealed the precipitation of coldest quarter contribute significant role in predicting the wolf habitat distribution between 3000m to 5300m. This study finds that, grassland with the slope less than 20 degree and area near to the river are best for potential distribution of wolves, and acts as the baseline information for further research and conservation despite of very few studies and information documented in the Nepalese Himalaya. Imminent action and more studies should be initiated for species conservation. Furthermore, research on elusive animals like wolf, their suitable prey base and human-wolf relationship by building communal harmony and through effective conservation techniques is very important for future.
Thesis
Full-text available
Annapurna Conservation Area (ACA) is highly potential and domineering ecological habitats for wildlife and fauna. The region is home to elusive wild animals such as Himalayan wolf, Snow leopard and Tibetan fox. Himalayan Wolf (Canis lupus chanco) is the ancient linage of the dog-clade which is historically inhabiting in the Nepalese Himalaya, which has been extirpated from the most of its historic range since a couple of decades. Majority of the people in the region depends on livestock herding and tourism. Maximum Entropy (MAXENT) model was used to find the potential habitat of Himalayan wolf in Upper mustang, ACA. Presence-only data were collected throughout the feasible 2*2 Km2 grids. To get the better result 13 environmental variables were computed. Raster images were worked in ArcGIS and EDRAS Imagine and computed in MAXENT. The Naïve occupancy of Himalayan wolf was estimated 71.87%. Relative Availability Index and average entropy values were 2 sign samples/ km and 7.929 respectively predicting the whole study area potential wolf habitat. 1089 km2 area is estimated as potential Himalayan wolf habitat out of 2076.56 km2 studied. The study also shows high habitat overlapping between carnivores and livestock. MAXENT model, has shown high level of predictive performance with an ‘Area Under Curve’ between 0.876±0.983 and average Standard Deviation 0.044. It is understood that AUC value closer to 1, more suitable the model against computed predictive variables whereas lesser the value of Standard Deviation, less chance of overfitting among the presence data. The model has revealed the precipitation of coldest quarter contribute significant role in predicting the wolf habitat distribution between 3000m to 5300m. This study finds that, grassland with the slope less than 20 degree and area near to the river are best for potential distribution of wolves, and acts as the baseline information for further research and conservation despite of very few studies and information documented in the Nepalese Himalaya. Imminent action and more studies should be initiated for species conservation. Furthermore, research on elusive animals like wolf, their suitable prey base and human-wolf relationship by building communal harmony and through effective conservation techniques is very important for future.
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The origin and evolution of the domestic dog remains a controversial question for the scientific community, with basic aspects such as the place and date of origin, and the number of times dogs were domesticated, open to dispute. Using whole genome sequences from a total of 58 canids (12 gray wolves, 27 primitive dogs from Asia and Africa, and a collection of 19 diverse breeds from across the world), we find that dogs from southern East Asia have significantly higher genetic diversity compared to other populations, and are the most basal group relating to gray wolves, indicating an ancient origin of domestic dogs in southern East Asia 33 000 years ago. Around 15 000 years ago, a subset of ancestral dogs started migrating to the Middle East, Africa and Europe, arriving in Europe at about 10 000 years ago. One of the out of Asia lineages also migrated back to the east, creating a series of admixed populations with the endemic Asian lineages in northern China before migrating to the New World. For the first time, our study unravels an extraordinary journey that the domestic dog has traveled on earth.Cell Research advance online publication 15 December 2015; doi:10.1038/cr.2015.147.
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The domestication of plants and animals over the past 11,500 years has had a significant effect not just on the domesticated taxa but also on human evolution and on the biosphere as a whole. Decades of research into the geo-graphical and chronological origins of domestic animals have led to a general understanding of the pattern and process of domestication, though a num-ber of significant questions remain unresolved. Here, building upon recent theoretical advances regarding the different pathways animals followed to become domesticated, we present a large-scale synthesis that addresses the global pattern of animal domestication alongside a discussion of the differ-ential evolutionary processes that have shaped domestic animal populations. More specifically, we present a framework for understanding how uncon-scious selection characterized the earliest steps of animal domestication and the role of introgression and the importance of relaxed and positive selection in shaping modern domestic phenotypes and genomes. 115
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The Qilian Mountain is located in the northeastern margin of Qinghai-Tibet Plateau. The authors had conducted two surveys on distributions of mammals in the Qilian Mountain and its adjacent area during the summer in 2001 - 2002. Altogether 72 species of mammals have been recorded in our investigational area, belongings to 7 orders, 20 families and 53 genera. Among them, 62 species which occupying 86.1% of the total number of the mammals are considered to be Palaearctic and 7 species to be Oriental mammals, besides, the others to be widely spread mammals. On the basis of comprehensive physical factors including altitude, landforms, climate, vegetation, hydrology, land-use, etc., the reasearch area was divided into 10 basic units (Operational Taxonomic Unit, OTU). Then the information of mammals in each unit was used to compute species composition similarity for the 10 units, using clustering methods. The clustering results indicated that the fauna on mammals in Huangshui Valley is quite different from all the other areas. The central area of Qilian Mountain is similar to the mountain area of Northern Qinghai Lake. The faunal differences between the Western Chaidamu Basin and the Mountain area of eastern Chaidamu Basin are obviously existent. The Mountain area of eastern Chaidamu Basin has a much closer relationship with the Central area of Qilian Mountain than that with the western and central area of Chaidamu Basin. Ecologically the existence of Qilian Mountain have caused obvious edge effects on mammal diversity and zoogeography. The species composition of mammals in this area not only have some similarity to Qinghai-Tibet Plateau, but also have a close relationship with Xinjiang area. For these reasons, the mammal diversity and zoogeography of the Qilian Mountain could be considered as a transitional area.
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From 2003 to 2004, the mammal fauna of the Jiuzhaigou National Nature Reserve was surveyed. Six separate surveys were conducted during this period. A summary of these collections showed that there are 78 species mammals in the Jiuzhaigou National Nature Reserve. An investigation of the biogeographic origin of this mammal fauna indicates that 50 species belong to the Oriental realm, 25 species belong to the Palaearctic realm, and 3 species are widely distributed. Ten general patterns of distribution were noted among the 78 species of mammal. This reserve's mammals show a very high south-north penetrability and antiquity. Twenty species in this reserve have been identified as nationally protected rare mammals; six of the species are first grade protected rare mammals. In addition, there are many rare small mammals in the reserve, such as the China jumping mouse (Eozapus setchuanus), Chinese Dormouse (Chaetocaudo sichuanensis) and Bedford's vole (Proedromys bedfordi). There is a distinct trend for an increasing proportion of Palaearctic mammals and decreasing proportion of Oriental mammals as one ascends from 2 000 - 3 600 m. However, 3 200 - 3 600 m there is an abundance of both Palaearctic and Oriental species. These results show that this mammal fauna is broadly in transition in this area, including at elevations above 3 200 m.
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The golden jackal of Africa (Canis aureus) has long been considered a conspecific of jackals distributed throughout Eurasia, with the nearest source populations in the Middle East. However, two recent reports found that mitochondrial haplotypes of some African golden jackals aligned more closely to gray wolves (Canis lupus) [1, 2], which is surprising given the absence of gray wolves in Africa and the phenotypic divergence between the two species. Moreover, these results imply the existence of a previously unrecognized phylogenetically distinct species despite a long history of taxonomic work on African canids. To test the distinct-species hypothesis and understand the evolutionary history that would account for this puzzling result, we analyzed extensive genomic data including mitochondrial genome sequences, sequences from 20 autosomal loci (17 introns and 3 exon segments), microsatellite loci, X- and Y-linked zinc-finger protein gene (ZFX and ZFY) sequences, and whole-genome nuclear sequences in African and Eurasian golden jackals and gray wolves. Our results provide consistent and robust evidence that populations of golden jackals from Africa and Eurasia represent distinct monophyletic lineages separated for more than one million years, sufficient to merit formal recognition as different species: C. anthus (African golden wolf) and C. aureus (Eurasian golden jackal). Using morphologic data, we demonstrate a striking morphologic similarity between East African and Eurasian golden jackals, suggesting parallelism, which may have misled taxonomists and likely reflects uniquely intense interspecific competition in the East African carnivore guild. Our study shows how ecology can confound taxonomy if interspecific competition constrains size diversification. Copyright © 2015 Elsevier Ltd. All rights reserved.
Article
We explored multiple linkages among grey wolves (Canis lupus), elk (Cervus elaphus), berry-producing shrubs and grizzly bears (Ursus arctos) in Yellowstone National Park. We hypothesized competition between elk and grizzly bears whereby, in the absence of wolves, increases in elk numbers would increase browsing on berry-producing shrubs and decrease fruit availability to grizzly bears. After wolves were reintroduced and with a reduced elk population, we hypothesized there would be an increase in the establishment of berry-producing shrubs, such as serviceberry (Amelanchier alnifolia), which is a major berry-producing plant. We also hypothesized that the percentage fruit in the grizzly bear diet would be greater after than before wolf reintroduction. We compared the frequency of fruit in grizzly bear scats to elk densities prior to wolf reintroduction during a time of increasing elk densities (1968-1987). For a period after wolf reintroduction, we calculated the percentage fruit in grizzly bear scat by month based on scats collected in 2007-2009 (n = 778 scats) and compared these results to scat data collected before wolf reintroduction. Additionally, we developed an age structure for serviceberry showing the origination year of stems in a northern range study area. We found that over a 19-year period, the percentage frequency of fruit in the grizzly diet (6231 scats) was inversely correlated (P < 0·001) with elk population size. The average percentage fruit in grizzly bear scats was higher after wolf reintroduction in July (0·3% vs. 5·9%) and August (7·8% vs. 14·6%) than before. All measured serviceberry stems accessible to ungulates originated since wolf reintroduction, while protected serviceberry growing in a nearby ungulate exclosure originated both before and after wolf reintroduction. Moreover, in recent years, browsing of serviceberry outside of the exclosure decreased while their heights increased. Overall, these results are consistent with a trophic cascade involving increased predation by wolves and other large carnivores on elk, a reduced and redistributed elk population, decreased herbivory and increased production of plant-based foods that may aid threatened grizzly bears.