Abstract
Wildlife conservation in the Anthropocene requires bold conservation solutions including restoration of ecosystems and species. The recovery of large carnivore populations is a conservation goal which can generate significant benefits in terms of ecosystem services, ecological functionality, and human well-being. Tigers Panthera tigris, Asia’s most iconic species, are currently restricted to less than 10% of their historic range with recent national extinctions from a number of countries in mainland Southeast Asia. Tiger recovery through range expansion requires suitable habitat, a robust prey base, and high levels of institutional support for conservation. We explored government support for conservation to produce a ranking of the political opportunities for tiger restoration across current and former tiger range countries. We used this analysis, in combination with globally remotely sensed data-sets on human impact, to show that there is potential for significant tiger range expansion. We identified large expanses of currently unoccupied, but potentially suitable, habitat in at least 14 countries including all extant tiger range countries and four countries with extirpated tiger populations – Cambodia, Lao PDR, Viet Nam, and Kazakhstan. Thirty-two percent of expansion areas were within 50-km, and 50% within 100-km, of current tiger populations highlighting that in many landscapes range expansion could be driven by the natural dispersal of tigers provided connectivity is maintained or enhanced. The proportion of potential range within existing protected areas varied between <5% in India, Indonesia, and China, to >60% in Thailand and Cambodia. As such socially appropriate conservation approaches, in collaboration with local communities, will be necessary to support tiger recovery in many areas. We recommend that some of the areas which we have identified should be highlighted as significant for future tiger conservation by tiger range country governments. Whilst the landscapes and sites which we identify will require detailed ground-truthing, and all tiger reintroductions need extensive planning and feasibility assessments, safeguarding these areas for human-carnivore coexistence could provide significant planetary benefits and support both tiger recovery and Global Sustainable Development Goals.
1 Introduction
Wildlife conservation in the Anthropocene requires bold conservation solutions. Current global conservation efforts have largely failed, and more ambitious commitments and innovations are required to stem wildlife declines (; ). Such innovations need to move beyond protection to the re-expansion of nature including ecosystem restoration and rewilding (). The UN Decade on Ecosystem Restoration (2021-2030) is a rallying call for the protection and revival of ecosystems and highlights that a strong connection exists between recovering nature and sustainable human development (). Large carnivores have the potential to be leveraged as symbols for ecosystem restoration. They are amongst the most charismatic and ecologically significant animals and require large expanses of suitably managed habitat (; ). Restoring, then maintaining, viable wild populations of large carnivores, can act as a driver to preserve larger, better connected, and better-quality ecosystems. The majority of the world’s terrestrial large carnivores are restricted to a fraction of their historic ranges and restoring carnivore distribution is a conservation goal which can also generate significant benefits in terms of ecosystem services, ecological functionality, and human well-being (; ). The Anthropocene extinction crisis is particularly acute in Asia and is exemplified by the state of the continent’s most iconic species: tiger Panthera tigris. Whilst global tiger populations, if not distribution, are increasing, tigers remain the world’s most threatened large cat. There are fewer than 5,000 wild tiger individuals and these are restricted to less than 10% of their historic distribution (; ). Since ~1850, tigers have been lost from at least 14 countries with three national extirpations, in Viet Nam, Lao PDR, and Cambodia, having occurred in the past 25 years (; ; ). The successful long-term recovery of tigers requires both securing current source populations () and expanding the species’ occupied range. Under the recently developed IUCN Green Status Assessment () tiger likely meets the criteria for Critically Depleted. Increasing the species’ distribution, and the ecological breadth of places where tiger occur and are functional, is necessary to recover the species globally. Implementing actions to reverse the centuries-long decline in tiger range is an ambitious and politically relevant conservation goal. Such planning needs to be long-term and could help create proactive and inspirational conservation goals which move beyond defending current tiger space and allow tiger populations, and conservation successes, to expand.
Increases in tiger range can be driven by both natural range expansion, through dispersal from current tiger population sources, as well as planned translocations and reintroductions into parts of the historic distribution from which tigers have been lost (; ). Both of these processes are known to have driven large carnivore recovery in Europe (). Natural dispersal of grey wolves Canis lupus from sources in remote and mountainous strongholds, combined with specific reintroduction and translocation programs for Eurasian lynx Lynx lynx and brown bear Ursus arctos, have resulted in significant range expansion of these carnivores in recent decades (). Strong legislative frameworks and political support for law enforcement, combined with increasing social tolerance for carnivores and rural depopulation, were significant drivers of this range expansion across Europe (; ). Despite their appeal to many people, often distant from carnivore occupied landscapes, the conservation and recovery of large carnivores can be controversial (; ; ). Large carnivores can compete with people for space and resources. Human wildlife conflict, both real and perceived, can impact attitudes to carnivore recovery globally (; ). Most successful examples of carnivore range recovery occur in countries and landscapes with strong governmental policies facilitating and resourcing conservation and with support from local communities (). Similarly, landscapes with increasing tiger populations are often characterised by high levels of law enforcement, good management of species and their habitats, high community support, and local economies and jobs created around wildlife (e.g. tourism) (; ). However, when assessing where the conditions for tiger recovery are present, less attention has been paid to the enabling political conditions which may support tiger conservation.
We explored issues linked to government support for conservation to produce a ranking of the political opportunities for tiger restoration across 30 current and former tiger range countries. We used this analysis, in combination with globally remotely sensed data-sets on human impact, to identify possible opportunities for tiger range expansion across Asia. We compared opportunities for tiger range expansion with current conservation priorities and focuses, measured through overlap with government protected areas and Key Biodiversity Areas (KBAs; ). Identifying range expansion opportunities and constraints is important to guide the future tiger conservation agenda and to proactively identify spaces for possible future tiger conservation. Landscapes and sites with opportunities for tiger range expansion are likely to require conservation interventions at all levels from national government to local communities. Protecting such future tiger space, in collaboration with local stakeholders, will ensure conservationists remain ahead of the curve on global tiger recovery.
2 Materials and methods
We mapped the historic range of tiger through identifying terrestrial ecoregions from which tiger were reported based on georeferenced historic tiger records, indicative of a breeding population, from between ~1750 and 2020. This historic resident breeding distribution (henceforth historic distribution) covered 11,792,218-km2 in 30 countries (Figure 1). We mapped the current tiger distribution (henceforth current distribution) based on ‘Extant’ areas within the 2015 IUCN Red List assessment of tiger () which we modified to account for recently documented extirpations (; ). This current tiger distribution covered 673,737 km2 in 10 countries (Figure 1; SI Table 1). Multiple factors influence distribution of tigers including habitat structure and, notably, prey abundance (; ). However, carnivore distribution may also be influenced by human pressures and behaviour which could be manifested in, for example, levels of retaliatory killing of tigers or elevated hunting pressures on prey species. We hypothesised that current (and future) tiger distribution is strongly influenced by human pressures and that the relationship between human pressure and probability of tiger presence differs between regions due to political, cultural, and ecological factors (; ). We used the global Human Modification Index (HMI) to establish the relationship between human impact and current tiger presence. HMI is a global 1-km2 resolution raster data-set indicating the impact of human activity and comprising data on human settlement, agriculture, transportation, mining and energy production, and electrical infrastructure (). We calculated the mean (plus-minus Standard Deviation) HMI score of polygons within current tiger distribution in each of the ten countries in which tiger currently occur and within each of three continental regions (South Asia; Southeast Asia; East-Central-West Asia).
Figure 1
Within each country in which tiger currently occur we identified polygons of area greater than 500 km2, equivalent to the size of the smaller Tiger Conservation Landscapes identified by
In addition to the direct human impact, indicated by the HMI score, the appropriateness of countries and landscapes for large carnivore recovery may be dependent upon a supportive political environment. We scored each of the 30 countries which comprised the historic tiger distribution based on perceived political support for conservation. Data we used provided information on general support to biodiversity conservation and specific information on planning for carnivore conservation. We collated data on domestic conservation funding (
To identify possible landscapes for tiger range expansion we combined the HMI polygons with the political support scores. For countries identified as likely highly supportive of large carnivore conservation all polygons with a mean HMI value smaller (i.e. less human impact) than the mean plus one standard deviation of occupied tiger polygons for the respective country or continental region (for countries from which tiger have been extirpated) were selected. For countries with intermediate levels of political support all polygons with HMI values less than the mean value for occupied tiger landscapes were selected, and for those with low support all polygons with HMI less than the mean value minus one standard deviation were selected. To identify overlap between current conservation priorities and possible tiger range expansion areas we compared range expansion polygons with current protected area coverage from the World Database of Protected Areas and the locations of KBAs (
3 Results
Our mapped current tiger distribution covers ~674,000 km2 in ten countries (SI Table 1; Figure 1). This represents 5.7% of the historic tiger distribution (Figure 1). The mean Human Modification Index (HMI) within current tiger distribution varied among the ten extant tiger range countries (SI Table 1; Figure 2) and was highest in India, Bangladesh, and Nepal and lowest in Russia and Myanmar. The HMI of the current tiger distribution was higher in South Asia than in Southeast and East Asia, and higher in Southeast Asia than East Asia (SI Table 1). Almost three-quarters of current tiger distribution was classified as forest (SI Table 1). Lowest forest cover of current tiger distribution was in Nepal and India, where 40 and 55% of current tiger distribution respectively, was in human modified habitats (SI Table 1).
Figure 2

Mean (± Standard Deviation) Human Modification Index (HMI) scores for current tiger distribution in each extant tiger range country.
3.1 Political support indicators
Domestic conservation funding across the 30 current and former tiger range countries varied between 0 (10 countries) and 82 million USD/year (South Korea) with a mean of 13.7 million USD/year for the period for which data was available i.e. 2001-2008 (
3.2 Mapping areas for tiger range expansion
The HMI values we used to identify areas for possible tiger range expansion varied from 0.52 in India, to <0.01 for the six former tiger range countries with low levels of political support for conservation in Central and West Asia (SI Table 4). Using these thresholds, we identified 1,293,921-km2 within 176 blocks of habitat ( 7,346-km2; range 500 – 502,007-km2) across the historic tiger distribution that is potentially suitable for range expansion (Figure 1). These areas (henceforth ‘expansion areas’) occurred across 14 countries (Table 1) including all extant tiger range countries (92.7% of predicted expansion area) and four countries with extirpated tiger populations – Cambodia, Lao PDR, Viet Nam, and Kazakhstan. The countries with the largest extent of expansion areas were India (612,718-km2), China (201,656-km2), and Russia (137,684-km2). These countries comprised >70% of all expansion areas. No expansion areas were identified from 16 countries within the historic tiger distribution including North and South Korea and the majority of countries in central and western Asia. In total 32% of expansion areas were within 50-km of current tiger distribution and 50% were within 100-km. Four countries (Indonesia, Bangladesh, Malaysia, and Bhutan) had >50% of their expansion areas within 50-km of current tiger distribution (Table 1; SI Table 5).
Table 1
| Country | Area (km2) | Number of blocks | % within 50-km | % within 100-km | % within Protected Areas | % Forest |
|---|---|---|---|---|---|---|
| India | 612,718 | 93 | 44.2 | 71.5 | 4.6 | 25.1 |
| China | 201,656 | 26 | 1.9 | 7.3 | 3.1 1 | 63.8 |
| Russia | 137,684 | 10 | 27.8 | 34.6 | 16.2 | 96.7 |
| Indonesia | 114,792 | 12 | 72.5 | 90.5 | 3.6 | 43.2 |
| Thailand | 94,965 | 19 | 9.3 | 24.3 | 63.9 | 92.9 |
| Cambodia | 70,868 | 2 | 0.0 | 0 | 78.3 | 90.0 |
| Myanmar | 23,688 | 2 | 26.8 | 65.9 | 17.2 | 97.0 |
| Vietnam | 14,501 | 2 | 0.0 | 0 | 18.6 | 95.2 |
| Kazakhstan | 8,617 | 3 | 0.0 | 0 | 30 | 0.1 |
| Bangladesh | 7,050 | 1 | 55.1 | 88.6 | 8.1 | 97.9 |
| Laos | 3,000 | 1 | 0.0 | 0 | 9.2 | 99.5 |
| Malaysia | 2,116 | 2 | 100.0 | 100 | 5.9 | 59.3 |
| Nepal | 1,524 | 2 | 35.2 | 35.2 | 28.9 | 87.4 |
| Bhutan | 742 | 1 | 100.0 | 100 | 18.5 | 99.4 |
Range expansion area (km2) per country plus the number of discrete blocks of habitat and the percentage of expansion area within 50-km and 100-km of current tiger distribution, the percentage within Protected Areas, and the percentage classified as Forest (SI Table 2).
1. Protected area coverage for China excludes the Northeast Tiger and Leopard National Park which is not yet included within the World Database of Protected Areas (www.protectedplanet.net).
The proportion of expansion areas within existing protected areas varied between <5% in India, Indonesia, and China to 64% in Thailand and 78% in Cambodia (Table 1). Overall, 188,066-km2 of expansion areas (14%) were within protected areas. Overlap between protected areas and range expansion areas was low in the four extant tiger range countries in South Asia (4.7%) and highest in mainland Southeast Asian (30%). Just under 14% of range expansion areas overlapped KBAs (SI Table 4). Overlap with KBAs was highest in Lao PDR and Myanmar (>60%) and lowest in Indonesia (4.8%) and Russia (3.4%). More than 500,000-km2 (41.5%) of range expansion area was in human modified habitat including large areas of low intensity agricultural and forest mosaics in India (SI Table 6). In Malaysia and Indonesia almost half of expansion area was in human modified habitats. In 8 of 14 countries, including all Southeast Asian countries apart from Malaysia and Indonesia, >90% of range expansion areas were in forest (SI Table 6).
4 Discussion
We demonstrate that there is potential for significant tiger range expansion across the species’ historic distribution with large expanses of currently unoccupied, but potentially suitable habitat, remaining in 14 countries. Whilst the global tiger population may be increasing from a nadir in the first decade of the 21st century, the species’ range contraction continues (
The expansion areas which we have identified are not a prescriptive blueprint for tiger recovery, reintroductions, or translocations. Instead, we hope that they represent some of the opportunities for future range expansion provided that the landscapes are protected, prey are sufficiently abundant, and threats are mitigated. Many of the sites identified may be worse on the ground than predicted – particularly in terms of tiger prey density, levels of effective land-use management, and community support. There is a need for country and landscape-specific ground-truthing of the expansion areas and global analyses cannot replace the need for detailed site-based assessments. Such studies, using more up-to-date and accurate information on the current distribution and status of tigers, will refine our analysis. Such country specific assessments are critical with regard to understanding habitat quality and tiger prey densities: important factors for tiger recovery but which cannot be obtained through remotely sensed data-sets. We hope that country specific assessments will be used to develop global goals for tiger range expansion which could be formalized as part of the revision of the Global Tiger Recovery Program (GTRP) for the next 12-years.
We demonstrate that the relationship between human activity, identified through the Human Modification Index (HMI), and the presence of tigers differs significantly between countries and regions. This relationship is likely driven by levels of anthropogenic mortality to tigers and tiger prey which in turn is linked to levels of tolerance for living with large mammals (both carnivores and ungulates) as well as the degree of enforcement of legal protection. The HMI of occupied tiger areas was higher in South Asia, particularly within India, Bangladesh and Nepal, than across the rest of the current tiger distribution. Understanding the political, cultural, and ecological mechanisms by which tigers are able to coexist with different intensities of human activity is essential for global tiger recovery. Combining HMI with political support scores adds an extra nuance to our identification of where future tiger conservation opportunities may exist. Whilst Asian countries have been identified generally as underperformers in megafauna conservation (
We identified three of the thirty indigenous tiger range countries as being highly supportive for large carnivore conservation: India, Thailand, and South Korea. These conclusions are validated by some independent information such as India’s success in tiger conservation (
In South and Southeast Asia, the majority of range expansion areas were within 100-km of current tiger distribution: significantly less than the documented straight-line dispersal distance of tigers (
For expansion areas that are isolated from current tiger distribution, reintroductions can be considered. Large carnivore reintroductions are an increasingly widespread, and increasingly successful, conservation tool. Tiger reintroductions using wild captured or rehabilitated tigers, have been successful in a number of tiger range countries including India and Russia (
Across much of the tiger’s historic distribution the density of tiger prey species is significantly depleted due to both legal and illegal hunting. Half of the mammalian prey species of tiger are threatened with extinction, and roughly 80% have decreasing population trends (
Given their large home-ranges the conservation of carnivores can compete for space with economic development, including extractive industries and infrastructure. Many areas of low human impact globally are targeted for exploitation particularly for minerals and fossil fuels.
As with all global conservation analysis our results are impacted by the quality of the data used. Neither our current or historic tiger distributions will be completely accurate. The current tiger distribution from
The St Petersburg Tiger Summit in 2010 and the range-wide endorsement of the Tx2 Goal have revolutionised tiger conservation and spurred unprecedented conservation efforts and investments (
Statements
Data availability statement
The original contributions presented in the study are included in the article/Supplementary Material. Further inquiries can be directed to the corresponding author.
Author contributions
TG, SC conceived the study. TG, RR, PI, DP conducted the analysis. TG, RR, GJ, PI, JY, LK, AL, MP, PC, QJ, WR, JR, SR, NS, CW wrote the manuscript. All authors contributed to the article and approved the submitted version.
Acknowledgments
Michael Belecky, Ashley Brooks, Smriti Dahal, Rajesh Gopal, Abigail Hehmeyer, Grigory Mazmaniants, and Rohit Singh provided advice and insights during the development and drafting of this paper. Paolo Tibaldeschi assisted with data analysis. WWF US, WWF Sweden, WWF Germany, WWF UK, and WWF International provided funding support for this work.
Conflict of interest
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
Publisher’s note
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.
Supplementary material
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fcosc.2023.1124340/full#supplementary-material
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Summary
Keywords
restoration, landscape, tiger conservation, Asia, carnivore, protected area management, reintroduction
Citation
Gray TNE, Rosenbaum R, Jiang G, Izquierdo P, Yongchao JIN, Kesaro L, Lyet A, Pasha MKS, Patterson DJ, Channa P, Jinzhe QI, Ripple WJ, Roberts JL, Roy S, Shwe NM, Wolf C and Chapman S (2023) Restoring Asia’s roar: Opportunities for tiger recovery across the historic range. Front. Conserv. Sci. 4:1124340. doi: 10.3389/fcosc.2023.1124340
Received
15 December 2022
Accepted
15 March 2023
Published
20 April 2023
Volume
4 - 2023
Edited by
Zuofu Xiang, Central South University Forestry and Technology, China
Reviewed by
Xueyou Li, Kunming Institute of Zoology (CAS), China; Chris Carbone, Zoological Society of London, United Kingdom
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Copyright
© 2023 Gray, Rosenbaum, Jiang, Izquierdo, Yongchao, Kesaro, Lyet, Pasha, Patterson, Channa, Jinzhe, Ripple, Roberts, Roy, Shwe, Wolf and Chapman.
This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Thomas NE Gray, tgray@wwf-tigers.org
This article was submitted to Animal Conservation, a section of the journal Frontiers in Conservation Science
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