Reconstructing the historical distribution of the Amur Leopard (Panthera pardus orientalis) in Northeast China based on historical records

Abstract

The range of the Amur leopard (Panthera pardus orientalis) has decreased dramatically over the last 100 years. This species is still under extreme risk of extinction and conservation efforts are rigorous. Understanding the long-term dynamics of the population decline would be helpful to offer insight into the mechanism behind the decline and endangerment and improve conservation perspectives and strategies. Historical data collection has been the challenge for reconstructing the historical distribution. In China, new gazetteers having systematic compilation and considerable local ecological data can be considered as an important complementary for reconstruction. Therefore, we have set up a data set (mainly based on the new gazetteers) in order to identify the historical range of the Amur Leopard from the 1950s to 2014. The result shows that the Amur leopard was historically widely distributed with large populations in Northeastern China, but it presented a sharp decline after the 1970s. The decline appeared from the plains to the mountains and northeast to southwest since the 1950s. Long-term historical data, mainly from new gazetteers, demonstrates that such resources are capable of tracking species change through time and offers an opportunity to reduce data shortage and enhance understanding in conservation.

Full text

Reconstructing the historical distribution of the Amur Leopard... 143
Reconstructing the historical distribution of the Amur
Leopard (Panthera pardus orientalis) in Northeast China
based on historical records
Li Yang1, Mujiao Huang1,*, Rui Zhang1,*, Jiang Lv1, Yueheng Ren1,
Zhe Jiang1, WeiZhang1, Xiaofeng Luan1
1 School of Nature Conservation, Beijing Forestry University, NO.35 Tsinghua East Road Haidian District,
Beijing, 100083, P. R. China
Corresponding author: Xiaofeng Luan (luanxiaofeng@bjfu.edu.cn)
Academic editor: E. Eizirik  |  Received 1 November 2015|Accepted 9 May 2016| Published 25 May 2016
http://zoobank.org/B6BF470C-504C-46DE-A4DD-4798C72CD4B1
Citation: Yang L, Huang M, Zhang R, Lv J, Ren Y, Jiang Z, Zhang W, Luan X (2016) Reconstructing the historical
distribution of the Amur Leopard (Panthera pardus orientalis) in Northeast China based on historical records. ZooKeys
592: 143–153. doi: 10.3897/zookeys.592.6912
Abstract
e range of the Amur leopard (Panthera pardus orientalis) has decreased dramatically over the last 100
years. is species is still under extreme risk of extinction and conservation eorts are rigorous. Under-
standing the long-term dynamics of the population decline would be helpful to oer insight into the
mechanism behind the decline and endangerment and improve conservation perspectives and strategies.
Historical data collection has been the challenge for reconstructing the historical distribution. In China,
new gazetteers having systematic compilation and considerable local ecological data can be considered as
an important complementary for reconstruction. erefore, we have set up a data set (mainly based on
the new gazetteers) in order to identify the historical range of the Amur Leopard from the 1950s to 2014.
e result shows that the Amur leopard was historically widely distributed with large populations in
Northeastern China, but it presented a sharp decline after the 1970s. e decline appeared from the
plains to the mountains and northeast to southwest since the 1950s. Long-term historical data, mainly
from new gazetteers, demonstrates that such resources are capable of tracking species change through
time and oers an opportunity to reduce data shortage and enhance understanding in conservation.
Keywords
Amur Leopard, Far Eastern Leopard, historical distribution, new gazetteers, Northeast China, Panthera
pardus orientalis
* e author contributed equally
ZooKeys 592: 143–153 (2016)
doi: 10.3897/zookeys.592.6912
http://zookeys.pensoft.net
Copyright Li Yang et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0),
which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
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Li Yang et al. / ZooKeys 592: 143–153 (2016)
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Introduction
Long-term historical data would oer insight into understanding the ecological and
biogeographic characteristics of population decline, and help develop predictive power
for conservation management (Boakes et al. 2010; Chapron et al. 2014; Rondinini and
Visconti 2015; Turvey et al. 2015). However, despite recognition of the considerable
potential of long-term datasets for conservation research, policy and practice, applying
long-term ecology data is still restricted by practical and conceptual barriers, includ-
ing data accessibility, spatially and temporally variable and non-standardized sampling
(Hortal et al. 2008; McClenachan et al. 2012; Davies et al. 2014). erefore, recent
studies always address time-scales of less than a decade (Davies et al. 2014). Only a few
researchers use records over periods longer than 20 years (Vandel and Stahl 2005; Pri-
gioni et al. 2007; Rick et al. 2012; Turvey et al. 2015). ere is an increasing awareness
of the need, not only to integrate historical data into conservation and environment
management, but also to assess the usefulness and potential limitation of this data for
developing our understanding of long-term species change.
Local gazetteers, called “difangzhi” (地方志), also translated as local records or
annals, contain abundant information on environmental conditions and resources in
China (Wen et al. 2006; Looney 2008). New gazetteers typically record considerable
local ecological data from the 1950s, including animal records, as well as economic,
political and demographic information (Looney 2008; Xue 2010). Most of them are
compiled following a specic “scientic” natural history tradition and this makes it
easy to identify the species records. In addition, compilation of gazetteers is systema-
tized and provides a geographical coverage across most of China. For the records
from eld surveys, museums and papers, new gazetteers may have been neglected
for a long time. is may be seen as reasonable, because new gazetteers are only
concerned with species which are economic, unique or common. Recent research,
however, highlights that this resource can provide eective records for reconstruct-
ing long-term population dynamics (Turvey et al. 2015; Zhang et al. 2016). In this
case, we suggest that new gazetteers can be an irreplaceable source for conservation
biology resources.
e Amur leopard, or Far Eastern leopard (Panthera pardus orientalis) is consid-
ered to be one of the most endangered subspecies in the world and has been listed
as critically endangered on the IUCN red list since 1996 (Uphyrkina et al. 2002).
Compared to the distribution in the late 19th century, the range of the Amur leopard
has decreased dramatically over the last 100 years. Recent research shows that there
are only 14–20 adults and 5–6 cubs in the southwestern Primorye region of Russia
(Henschel et al. 2008; Spitzen et al. 2012); Perhaps some may still occur in North
Korea, although their status there is uncertain (Uphyrkina et al. 2002); Fewer than 10
leopards likely existed in Jilin and Heilongjiang Provinces of northeast China in the
later 1990s, but the population seems to have increased in recent years (42 leopards
were photographed, including 40 adults (21 males, 17 females, and two of unknown
sex) and two cubs) (Yang et al. 1999; Wang et al. 2015). e tiny population that

Reconstructing the historical distribution of the Amur Leopard... 145
survives today is under extreme risk of extinction because of poaching, deforestation,
inbreeding, and anthropogenic pressure (Ma 1989; Yang et al. 1999; Yang et al. 2000;
Piao et al. 2011; Hebblewhite et al. 2011; Kelly et al. 2013; Cat Specialist Group
2014). erefore, a basic understanding of their distribution over decades is necessary
to design conservation strategies on a regional scale. To address this problem, we sug-
gest that new gazetteers can be a useful resource. Combining new gazetteers, which
regularly have Amur leopard records, with historical information from literature, news
and scientic surveys, we identied the range of the Amur leopard in Northeast China
from the 1950s to the 1990s. e historical dynamic for the Amur leopard in North-
east China was then reconstructed primitively by integrating historical records within
a geographic information system (GIS).
Materials and methods
Area
e study area comprises Heilongjiang province, Jilin province and northeast of
Inner Mongolia covered by temperate forest and boreal forest (N40°5'~53°17',
E115°30'~135°06', about 938000 km2 with a forested area of 402,000 km2), which is
the most important forestry and agricultural production base in China. e climate is
a continental monsoon climate with a negative water balance. e annual precipitation
is 400–1000mm, and the annual average temperature is 1–4°C, with the north - south
temperature gradient of 25. e region includes coniferous forest, broad-leaved mixed
forest, secondary forest, woodland shrub and marshy grass areas, and contains more
than 2500 plant species (Zhou 1997; Hou 2001). Coniferous forest is mainly located
in the Greater Khingan Mountains where it is characterized by Larix gmelinii. e
area of broad-leaved mixed forest includes the Lesser Khingan Mountains, Changbai
Mountains and Wanda Mountains dominated by Larix gmelinii, Pinus koraiensis and
Betula platyphylla (Cheng & Yan, 2008). Forest ecosystems in Northeast China sup-
port a number of forest-dependent carnivores and herbivores, for instance the Amur
leopard and its potential prey – the Siberian roe deer (Capreolus pygargus), wild boar
(Sus scrofa) and sika deer (Cervus nippon).
Data
Amur leopard historical records were obtained in the following ve ways:
(1) New gazetteers. Due to being an important cultural symbol, economic species
and having an ecological niche in human history and culture, large carnivores like
the Amur leopard are recorded regularly in new gazetteers. e data from new
gazetteers can be considered as the hard fact, because they came from the fur trade

Li Yang et al. / ZooKeys 592: 143–153 (2016)
146
records, hunter records, sightings, wild life surveys and conicts. Systematic com-
pilation made it possible to divide the records into dierent periods and to cover
most of China. Also utilizing fauna and nature reserve scientic surveys, papers,
scientic research and news, it is possible to compile substantial records for a long
period and covering a large geographical area. In this paper, more than 90% of his-
torical records came from new gazetteers. ey were obtained from the National
Library of China (http://www.nlc.gov.cn/), Duxiu Search Engine (http://www.
duxiu.com/), and the Wanfang database (http://www.wanfangdata.com.cn/). Rel-
evant information was found with the keywords – “Amur leopard”, “Far Eastern
Leopard” and “Panthera pardus orientalis”.
(2) Fauna surveys. Fauna surveys recorded the specimen data with geographic infor-
mation. We collected the distribution data from 9 Fauna surveys (Shou. 1962; Ma
et al. 1986; Gao et al. 1987; Ma et al. 1989; Wu. 1993; Wang. 1998; Wang et al.
1998; Sheng et al. 1999; Zhao et al. 1999; State Forestry Administration of the
People’s Republic of China 2009).
(3) Nature reserve scientic surveys. e relevant data was collected from the National
Library of China and the School of Nature Conservation, Beijing Forestry University.
(4) Papers and scientic research. e keyword “Amur leopard”, “Far Eastern Leop-
ard” and “Panthera pardus orientalis” were used to search several online databases
such as the China National Knowledge Internet, Wanfang Database, Duxiu Search
Engine, Google scholar (https://scholar.google.com/) and Biodiversity Heritage
Library (http://www.biodiversitylibrary.org/). Records from 1998 mainly came
from surveys (WCS program) in Heilongjiang and Jilin provinces.
(5) News. e keywords “Amur leopard”, “Far Eastern Leopard” and “Panthera pardus
orientalis” were used to search several online databases such as the China National
Knowledge Internet, Wanfang Database, Duxiu Search Engine, Science Daily and
Google. Records with detail location associated with photograph or video were
eective.
Data analysis
Filters are essential for further analysis, because historical records may contain poten-
tial errors or uncertainties. Records from four of the ways above (2-5) were integrated
into a dataset with brief information on the area of occupancy, and then this is com-
pared with the gazetteer records. Records that conicted with the dataset without any
hard facts, including relevant or detailed descriptions, were excluded from analysis. All
Chinese-language records were translated directly by the authors.
Maps of Northeast China (1:1,000,000) were acquired from the National Geo-
graphic Information Bureau, and spatial coordinates of all occurrence records were as-
signed using Google Earth. Multiple occurrences at matching sites, such as repetitions
of the same record or when the distance between locations was less than 5km were

Reconstructing the historical distribution of the Amur Leopard... 147
excluded. Records from killed animals, attacks on humans, prey remains, claw traces,
footprints and photographs were excluded if they were repeated. Valid records were as-
signed with coordinates from public resources such as Google Earth v 7.1.2, classied
into dierent decades (1950s, 1960s, 1970s, 1980s, 1990s and 2000–2014).
Amur leopard occurrence positions were then uploaded into ArcGIS 10.2, and
overlaid with layers representing altitude (http://srtm.csi.cgiar.org/) and the borders
of administrative regions. Using the number and intensity of records, it is possible
to reconstruct and evaluate the distribution change in dierent decades of the Amur
leopard population.
Results
A total of 84 documents of the 2235 new gazetteers examined presented information
on the Amur leopard. ese provided evidence regarding 294 Amur leopard records,
while another 169 Amur leopard records were retrieved from the other four sources
(as mentioned in the data sources section (2)-(5), giving a total of 463 Amur leopard
records). Some records were excluded (as mentioned in the methods section) and -
nally 341 Amur leopard records were mapped and used for further analysis (Table S1).
New gazetteers were considered as the main resource before the 1990s. In the 1950s,
96.4% of total records came from new gazetteers. 36.2% of the new gazetteers records
were conrmed by fauna and papers. 97.5%, 96.3% and 93.9% of total records came
from new gazetteers in the 1960s, 1970s and 1980s respectively. 42.4%, 41.2%, and
54.0% of the new gazetteers records were conrmed by fauna, papers and nature reserve
scientic surveys in the 1960s, 1970s and 1980s respectively. New gazetteers were still
an important resource in the 1990s (44.7% of total in the 1990s) (Table S1). Scientic
research for Amur leopard provided 17 records. Nature reserve scientic surveys and
news dominated after 2000 and only one record came from new gazetteers.
Our result implied that the Amur leopard was widely distributed with a large
population in Northeast China - Southern Lesser Khingan Mountains, Changbai
Mountain and Wanda mountains (Fig. 1). Before 1970, the Amur leopard occur-
rence points didn’t present obvious alterations (Fig. 1A, B). In the 1970s, the situ-
ation deteriorated continually. Compared with the distribution in the 1950s, there
was a reduction of 19.6% of records in the 1970s. From the 1980s onwards the Amur
leopard became limited to the core area of the Wanda Mountains and Changbai
Mountains. ere was a decline of 56.1% of records in the 1980s (Fig. 1C, D). In the
1990s, there were only a few records limited to several places, for example Hunchun,
Wanqing, Yanji and Helong. Afterwards, the distribution became concentrated in
the Hunchun and Wanqing in 2000–2014 (Fig. 1E, F). In addition, records in areas
higher than 200m continuously increased before 2000, but a slight decrease presented
in 2000–2014 (1950s: 79%; 1960s: 81%; 1970s: 84%; 1980s: 89%; 1990s: 92%;
2000–2014: 88%).

Li Yang et al. / ZooKeys 592: 143–153 (2016)
148
Figure 1. Amur leopard distribution in dierent periods.

Reconstructing the historical distribution of the Amur Leopard... 149
Discussion
Our investigation of the potential of using long-term historical records to reconstruct
long-term population dynamics shows that new gazetteers along with multiple eec-
tive resources can contribute novel insights for tracking the target species dynamics
across longer timescales than is usually addressed in ecology or conservation biology.
However, it is inevitable that the new gazetteer data compiled by non-scientic ob-
servers cannot provide a complete species records at the standard typically expected
by modern ecologists, even if combined with multiple resources (2-5 resources). For
example, hunt archives are the major component in new gazetteer records (like “Amur
leopard was killed by snare in Liushuihe Forest Farm in 1964, winter”). Since hunting
was banned in the 1980s, such records decreased sharply and can increase omission
error after 1990. Spatial and temporal bias in new gazetteer records also generates
deviation. For this reason, our investigation highlights some ways to reduce the com-
mission or omission errors in new gazetteer records, including choosing identiable
and economic species, no generic species absent in the same area, and integrating with
scientic data. Indeed, it is possible that the Amur leopard still exists in an area or
specic period with no record, or be present in contrary circumstances. However, such
historical dynamics for the Amur leopard in Northeast China still reveals some aspects
of the pattern and process of population decline across more than 50 years that can-
not be fully understood by the present-day population: e Amur leopard population
presents a sharp decline after the 1970s; and the decline is likely to appear from the
plain to the mountain and northeast to southwest, which implies negative impact from
anthropogenic pressure.
Some questions still need to be claried. For example, the Amur leopard popula-
tion in the Greater Khingan Mountains is elusive (the range is unknown, but they are
present). Only one fauna researcher (Gao et al. 1987) provides one record in Horqin
Right Front Banner, Inner Mongolia autonomous region in the southwestern Greater
Khingan Mountains. Four other fauna researchers describe the distribution briey
(Ma et al. 1986, 1989; Wu 1993; Zhao et al. 1999), and two of four suggest that local
population have been extinct in the 1970s with none appearing in hunting or witness
records after 1970 (Ma et al. 1989; Wu 1993). An extract taken from the Longjiang
xianzhi says” the leopard was bountiful at the time of the foundation of the PR of
China”. Longjiang town is about 80km away from the north of the Horqin Right
Front Banner. It is safe to say that the Amur leopard was distributed in the Greater Kh-
ingan Mountains in the 1950s, but the range change is still unclear. Moreover, higher
latitude records increased before 2000 and this implies that the negative impacts un-
der anthropogenic pressure for the Amur leopard and the decline in 2000-2014 can
be either a promising sign on the eectiveness of the conservation strategy, or such a
phenomenon may just be associated with data shortage.
Our reconstruction of the dynamics of the Amur leopard population decline en-
lightens a new case study with new perspectives shows that the historical records have
considerable potential to contribute to ecological baselines for informing conserva-

Li Yang et al. / ZooKeys 592: 143–153 (2016)
150
tionists. We recommend further investigation to not only detect feasibility for recon-
structing the dynamics of other species in China by Chinese gazetteers, or quantifying
the human pressure by the response of wildlife over time in modern China, but also
to evaluate the possibility of revealing the potential distribution in the past with the
historical records and distribution models. Furthermore, we encourage conservation
researchers and practitioners to consider this data type (not only the gazetteers, but also
other data resource considered as local historical record around the world) as comple-
mentary, in order to reduce data shortage and enhance understanding in conservation.
Sucient data would provide greater opportunities for conservationists and environ-
mental planners to put plans in place that would reduce the danger of endangered
species going extinct in China and elsewhere.
Acknowledgments
We thank Georey Pearce for improving language on this paper. is study was sup-
ported by a grant from the Ministry of Science and Technology of the People’s Re-
public of China (Research and application of key techniques on endangered species
conservation and prediction of forest re and pests in response to climate change,
2013BAC09B00).
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Reconstructing the historical distribution of the Amur Leopard... 153
Supplementary material 1
Distribution information from new gazetteers records and from other resources
Authors: Li Yang, Mujiao Huang, Rui Zhang, Jiang Lv, Yueheng Ren, Zhe Jiang, Wei
Zhang, Xiaofeng Luan
Data type: PDF le
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.
Supplementary material 2
Records in dierent periods from new gazetteers records
Authors: Li Yang, Mujiao Huang, Rui Zhang, Jiang Lv, Yueheng Ren, Zhe Jiang, Wei
Zhang, Xiaofeng Luan
Data type: PDF le
Copyright notice: is dataset is made available under the Open Database License
(http://opendatacommons.org/licenses/odbl/1.0/). e Open Database License
(ODbL) is a license agreement intended to allow users to freely share, modify, and
use this Dataset while maintaining this same freedom for others, provided that the
original source and author(s) are credited.