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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.
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 its 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) is 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).
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 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 report, the 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, and there is no comprehensive description of the current distribution of wolves across China. Therefore, we here summarize 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.
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; EO). 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, 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 (200) described the subspecies/subtypes of wolf in China and reported that the wolf was 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 pressence 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).
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 2011and in the two southernmost 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.
In addition to the literature investigation, we made a survey of wolf skins in the archives of Shaanxi Institute of Zoology, Kunming Natural History Museum of Zoology and the National Zoological Museum of China (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.
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.
In this study, we showed that contrary to what is reported in many references in the western literature, the 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 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 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.
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 of China. Hereby we corrected an error in western literature, in which most sources stated that wolves are 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.
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