Recovery of tigers in India: Critical introspection and potential lessons

Abstract


| INTRODUC TI ON
At a time when global biodiversity faces one its worst crises due to human actions of habitat destruction, climate change, pollution and direct exploitation of species (Johnson et al., 2017), the lessons from any successful conservation program need to be widely shared so that its relevant features can be replicated elsewhere. The tiger Panthera tigris, an apex predator, that requires vast habitat to harbour viable populations, serves as an umbrella species for conserving Asia's forest systems. Tigers have declined globally due to direct hunting, prey depletion and habitat destruction (Wikramanayake et al., 2010). The continued illegal trade of tiger body parts and products driven by high demand in China and South East Asia threatens the species with extinction (Dinerstein et al., 2007). Taking cognizance of the dire situation facing wild tiger survival, world leaders and conservation practitioners met at St. Petersburg, Russia, in 2010 to discuss strategies for tiger recovery (Joshi et al., 2016). This event was a first in human history where country leaders met to discuss conservation of a species. The outcome was a Global Tiger Recovery Program, that outlined strategies that may be undertaken singly or jointly by range countries to increase tiger numbers from a global estimate of ~3,643 in the year 2010 to ~5,845 (Table S1 and data sources) by the year 2022 and to protect tiger habitats (Global Tiger Initiative, 2011). Without dwelling on the scientific merits of achieving such a target, and judging from the most recent status of tigers in range countries, global tiger numbers have increased to around 4,981 (4,232-5,781; Figure 1, Table S1). India leads amongst the few other countries (Bhutan, Nepal, Russia and Indonesia; Figure 1, Table S1) that have recovered wild tiger populations. Bangladesh has since corrected their baseline tiger population status using the 4. In the Anthropocene it is unlikely that tigers will survive without targeted conservation investments. Political commitment and resources can become available for conservation when people and tigers benefit simultaneously. Conscious balance by governments between development for rapid economic prosperity and longterm ecological security will ensure that wild tigers and their intact ecosystems will survive for future generations.

K E Y W O R D S
economic evaluation, global tiger population, incentivized voluntary village relocation, management effectiveness evaluation, reintroduction, tiger conservation plans, tiger reserves F I G U R E 1 (a) Recent estimates of tiger numbers in range countries (see Table S1 and data sources); (b) Progress towards achieving the targeted tiger population number by range countries as depicted in the Global Tiger Recovery Plan. India and Bhutan have overachieved their targets while Nepal and Russia have made progress. Bangladesh has registered severe deficit due to an incorrect population baseline in 2010 modern scientific approach of spatially explicit capture-mark-recapture using camera traps (Borchers & Efford, 2008) to 106 (83-130) (Dey et al., 2015).
Herein, we provide a brief historical perspective, major conservation challenges, and highlight science-based policy and management strategies that have turned the tide for wild tiger conservation in India. Lastly, we outline the future actions required for conserving tigers and their ecosystems in India and elsewhere.

| HIS TORI C AL , RELI G I OUS AND CULTUR AL PER S PEC TIVE
Wildlife and tigers have been an integral part of Indian history, culture and religions. The most ancient records of tigers are found in the cave paintings of central India dated between 100,000 and 30,000 BP (Badam & Sathe, 1991). The seals and artefacts recovered from the civilizations of the Indus Valley depict tigers with the most ancient deity Shiva as Pashupatinath or the lord of animals (~5,000 YBP; Fairservis, 1983). The first Protected Areas in the world were declared in India by the Buddhist king Ashoka as Abhaya Aranya or 'forests without fear' around 250 BCE (Rangarajan, 2005). Several wild animals and plants, especially tigers are revered in religions that evolved within the Indian sub-continent (Hinduism, Jainism and Buddhism). These Eastern religions' doctrine of humans as custodians of nature that contrasts with the Abrahamic religious beliefs that promote humans to have dominion over nature (White, 1967). This philosophical mindset along with the concept of Ahimsa endows the society with a high level of tolerance towards all life forms that extends event to predators like tigers. This mindset is still the primary factor responsible for coexistence between high-density human populations and wildlife in India. However, this attitude is also responsible for sustaining a large population of feral livestock, dogs, and cats which are becoming a major problem for wildlife conservation. It was only after the colonial occupation of India that systematic removal of wildlife, including that of tigers, was undertaken by the Government through bounties and state sponsored actions (Rangarajan, 2005).
Modern conservation era began much after India's independence in the early 1970s, with the enactment of the Wildlife (Protection) Act (1972) (WPA, 1972). At the behest of J. C. Daniel and renowned ornithologist Salim Ali from the Bombay Natural History Society, the then Prime Minister of India, Indira Gandhi, took personal interest in tiger conservation and with assistance from the then World Wildlife Fund launched Project Tiger in 1973 as a central Government sponsored scheme in nine Protected Areas of India. The lion was subsequently replaced by the tiger as the national animal of India. India's tiger population was down to around 1,800 at the commencement of Project Tiger from a pre-colonial estimate of about 40,000 tigers (Panwar, 1982). The initial success of Project Tiger evaluated by the pugmark census technique put the figure to around 3,500 tigers by 1990s. This success brought in complacency in protection since legal hunting was banned in India and domestic market for tiger trophies was almost extinguished. However, during this period due to rapid economic growth in China and South East Asia (Milanovic, 2011), demand for tiger body parts for traditional medicine increased substantially. Owing to this increased demand, and after depletion of local tiger populations in China and SE Asia to unprofitable illegal harvest levels, tiger populations in India became the target for poachers.
While India basked in the glory of Project Tiger (Panwar, 1982), tiger populations in India were steadily being decimated by demand driven poaching. In 1994In -1995 in the famous Ranthambore Tiger Reserve were reduced to a handful by poachers, but it was only after the local extinction of tigers at Sariska Tiger Reserve in 2005 and associated media coverage, that then Prime Minister, Dr. Manmohan Singh appointed a Tiger Task Force to address the tiger conservation crisis in India.

| TI G ER CON S ERVATI ON P OLI C Y
The Tiger Task Force took cognizance of problems that plagued tiger conservation in India, some of that are universal across the range of the tiger, while others were unique to India. Tiger Task Force suggested implementable remedies through policy and management interventions (Narain et al., 2005). After a thorough research, the Tiger Task Force pinpointed major problems as systematic institutional failure, failure in protection due to alienation of poor local communities, misreporting of tiger numbers (Karanth et al., 2003), need for inviolate space for conservation, isolation of small Protected Areas and need for habitat connectivity-a landscape approach to tiger conservation planning, amongst others. The major recommendations of the Tiger Task Force were a paradigm shift for (tiger) conservation from being an exclusion model that alienated local communities to an inclusive one, where the local community became satisfied partners in conservation efforts. These recommendations led to the amend- A landscape approach to management planning; with provisions for an inviolate core zone that houses a viable tiger source population, a buffer zone that may be a sink habitat, with permitted multiple uses and habitat corridors that connect tiger populations within a landscape, was mandated for all tiger reserves (Gopal et al., 2007). For making core areas free of human settlements and use (inviolate), a monetary package of INR one million (USD 13,300; a small fortune by Indian standards) was offered per adult in a family by Project Tiger as a package for incentivized voluntary relocation. A major lacuna that was identified by the Tiger Task Force was the mechanism and associated procedural delays of fund transfer from the central Government (Project Tiger) to the tiger reserves. To address this, a local repository of funds for each tiger reserve was created in the form of a Tiger Conservation Foundation. Profits emanating from tourism activities based on tigers were to be shared with buffer zone communities by the park management as well as by the tourism industry. A monitoring protocol for tigers, associated fauna and their habitat, that was scientifically robust yet practical for large scale surveys was designed, tested and recommended to be implemented every four years across India and every year for all tiger reserves (Narain et al., 2005).

| Tiger reserves: Core, buffer and corridors
Currently, tigers occupy around 89,000 km 2 in India while forest area (potential habitat) within tigers' range is around 381,000 km 2 (Jhala et al., 2020). Thus, tigers were clearly limited by direct persecution and quality of their habitat (prey depletion). Of the total population of around 3,000 tigers in India 65% were within tiger reserves (Jhala et al., 2020). Although habitat was the least limiting factor yet, species like tigers can only thrive in legally protected human-free space. Such space in the form of core areas of tiger reserves is the most difficult requirement to secure in a densely populated, rapidly growing country like India. India has striven to increase areas gazetted as tiger reserves (Table S2).
The current 50 tiger reserves cumulatively protect an area of 72,810 km 2 of which, 40,145 km 2 is inviolate core. The average size of tiger reserves was 1,456 km 2 but varied between 492 (Orang TR) and 5,907 km 2 (Amrabad and Nagarjunsagar Srisailam tiger reserves combined as they are adjacent). Core areas across 50 tiger reserves averaged at 803 km 2 and varied between 79 and 4,762 km 2 (Table S2). The principle was to establish source populations (in core areas of tiger reserve) within larger sink habitats (buffer zones of tiger reserves and forested habitats across the landscape) that would have low-density tiger occupancy. A demographically viable tiger population requires a minimum of ~20 breeding females which translates to a population of between 75 and 100 tigers (Bisht et al., 2019;Chapron et al., 2008;Gopal et al., 2007). In India, due to the possibility of achieving high ungulate density, an average breeding tigress could successfully rear cubs within a territory of 40-50 km 2 . In some high prey density areas territories as small as 10-20 km 2 have been recorded for breeding tigresses (Y. Jhala & Q. Qureshi, unpubl. data;Sharma et al., 2010). A core area of 800-1,000 km 2 of a tiger reserve is the minimum required size to establish a source population. Many tiger reserves (42%) had core areas that could not potentially sustain 20 breeding females. Such tiger populations would remain viable only if they remained connected through habitat corridors as a metapopulation of two or more tiger reserves (Hanski et al., 1996). Tiger occupancy probability (Ψ) from across India was estimated through replicate ground surveys at every ~15 km 2 of forest patch and modelled using habitat, prey and human footprint indices. Habitat resistance to tiger movement was estimated as 1 − Ψ and corridors of least resistance between tiger reserves (protected areas) modelled using circuit theory in CIRCUITSCAPE (McRae et al., 2013). In central India, tiger movement across these corridors using non-invasive genetic sampling from 169 individual tigers was validated (Yumnam et al., 2014). All major habitat corridors connecting tiger populations within each landscape were modelled, mapped (Qureshi et

| Operationalizing inclusiveness of communities
Forests and even many Protected Areas in India are inhabited by people who depend on forest resources to a large extent to eke out a living (Wani & Kothari, 2007). With increasing human population such a lifestyle is unsustainable and forests continue to degrade due to overuse. Communities living within Protected Ares are unable to readily access many basic modern amenities like hospitals, school and higher education, and markets for purchase and selling their produce. Their livelihoods are threatened by depredation of livestock by large carnivores and crops raided by wild ungulates.
Therefore, when such communities are offered a genuine alternative, they are generally willing to resettle outside of the forest.
Unfortunately, during the initial years of Project Tiger, forest dwellers from within tiger reserves were evicted forcibly and without just compensation or handholding with alternative/new livelihoods (Wani & Kothari, 2007). These bitter experiences combined with strict restrictions on resource use and extraction from within tiger reserves by managers had alienated communities living within and on the edges of reserves. These disgruntled evictees readily assisted or themselves indulged in poaching tigers. In the experience of several of us, without the assistance of local communities, it is difficult to poach tigers.
The reforms that changed the relationship between communities and tiger reserves were:

Sharing of revenues generated from tiger reserve gate receipts
with communities residing in the buffer zone. The Tiger Task Force recommended that revenues generated from the tiger reserve by the Government as well as by the tourist industry should be shared with communities residing in the buffer zone. The prior practice was that all gate receipts were deposited with the State treasury and unavailable to the tiger reserve for its management.

To circumvent this limitation Tiger Conservation Foundations
were created for tiger reserves. Gate receipts, donations, funds for antipoaching activities (secret funds) could now be deposited with the tiger conservation foundation that were administered by a local committee with the Field Director of the tiger reserve as member secretary and representatives of local communities as committee members. Currently tiger conservation foundations are functional in 46 tiger reserves out of the 50 tiger reserves, F I G U R E 2 Tiger populations and density across occupied forests overlaid with habitat corridors connecting tiger reserves within each landscape (source: Jhala et al., 2020;Qureshi et al., 2014) with the past five-year average annual income of each tiger conservation foundation amounting to 287,000 (SE 75,000) USD.
Of this, on an average 40% of revenues generated by the tiger conservation foundations were to be used for buffer zone community welfare activities. Another 40% were used by the tiger reserve for enhancing protection. Tiger conservation foundations of all tiger reserves were not equally resourceful as some worldrenowned tiger reserves attracted more funds (e.g. Ranthambore Tiger Reserve averages at 2.5 million USD income per year) compared to other tiger reserves. At the discretion of the State Chief Wildlife Warden and the tiger conservation foundation committees the surplus funds from better-resourced tiger conservation foundations could be used for activities in the buffer zones of other tiger reserves within the State. These additional resources directly emanating from tiger reserves to communities provide the required incentives to support the conservation initiatives and convert hostile neighbours into supporters.

| Improving protection
The Wildlife (Protection) Act 1972 is a very powerful legislation. It provides for penalties of 3-7 years of imprisonment and/or fine of (c) Use of technology such as the E-eye, a sensor-based system of thermal, and visual cameras deployed along some sensitive borders of high-profile tiger reserves relayed real-time surveillance to tiger reserve managers and acted as major deterrent for poachers.
In some tiger reserves surveillance was also done with drones.

| Mitigating conflict
Ideally, it would be best not to mix humans and wildlife (especially large carnivores like tigers) through zoning (Linnell et al., 2012).
However, in densely populated countries like India availability of sufficient exclusive space to maintain viable populations of large carnivores is difficult and coexistence becomes an essential strategy for effective conservation (Woodroffe et al., 2005(Woodroffe et al., , 2014. The only exclusive space available for the global population of Asiatic lions is a 250 km 2 Gir-National Park, that is of insufficient size to hold a viable lion population. Around 500 lions coexist with humans in >10,000 km 2 of Saurashtra's agro-pastoral-industrial landscape .
Conflict is inevitable when humans and wildlife mix; crop raiding by wild ungulates, predation on livestock, attacks on humans and human deaths by large carnivores are major concerns which need to be effectively managed to prevent retaliation .
Although most States in Indian pay compensation for crop, livestock and human losses to wildlife, these were not free from corruption nor fair in relation to market prices of crops and livestock (Banerjee et al., 2013;Johnson et al., 2018). More recently attempts have been made to dispense compensation that is fair, transparent and corruption-free using mobile phone application-based documentation for authentication of claims and tracing of document processing with direct payment to the recipient's bank account. A quick, handsome monetary compensation dispensed with sensitivity helps ameliorate immediate anger and retaliation.
Compensation for human deaths ranged between 200,000 and 1,000,000 INR as Project Tiger compensation funds were often supplemented with state funds. The highest compensation was paid by the state of Maharashtra at one million INR for a human death. No amount of money can compensate loss of human lives, yet, very high compensation amounts for human deaths can lead to complacency in traditional precautionary systems and false claims of deaths attributed to tigers, especially amongst poverty riddled populations (Jhala & Sharma, 1997).

Tigers were responsible for an average of 38 (SE 4) human deaths
each year for the past 10 years in India. Tigers in the Sundarban tiger reserve (25%), from the Vidharbha region of Maharashtra (25%) and from around Pilibhit-Dudhwa tiger reserves (20%) accounted for 70% of all human deaths. Eleven tigers were eliminated (shot) in eight years (1.4, SE 0.46 tigers year −1 ) from the wild for being threat to human lives. As per the standard operating procedure (SOP) prescribed by NTCA (https://ntca.gov.in/ docum ents/#sop1) a tiger is to be removed from the wild (preferably captured) if it was found responsible for deliberate and consistent human attacks. Most attacks on humans were accidental when humans ventured into tiger habitat in search of non-timber forest produce. However, some tigers did become man eaters and it was only these individuals that were eliminated. A rapid response team constituted by a trained veterinarian and field staff, equipped with safe and rapidly acting anaesthetic drugs, good darting equipment, medical emergency kit and four-wheel drive vehicle was deployed at most tiger reserves or at the State level covering multiple tiger populations to address human-tiger (wildlife) conflict situations. With the advent of camera traps identifying problem tigers with some level of certainty has become easier. Nowhere else in the world is an individual animal involved in an attack on a human (be it accidental, provoked or deliberate) given a second chance. This was possible in India, perhaps due to the high tolerance level amongst the people due to cultural and religious reverence towards tigers. However, with tiger density increasing in multiple use areas, conflicts are on the increase and it may soon become necessary to implement population management either through capture and translocation or through reproductive control. Site-specific strategies would be required for tigers of the Vidharbha landscape and for tigers inhabiting sugarcane plantations around Pilibhit tiger reserve.
These populations pose exemplary problems that need planned mitigation for managing conflict caused by locally over abundant tigers. Such problems are likely to increase as conservation measures become more successful within reserves, and there is a scope of increasing tiger density in the surrounding human dominated landscapes due cropping patterns that provide cover and domestic livestock that provide alternate prey.

| Reintroductions and supplementation
In 2005 and 2009 India witnessed two local extinctions of tigers in its prestigious Sariska and Panna tiger reserves due to poaching.
Subsequently, two (one male and female) tigers were initially reintroduced to Sariska in 2008, and three (one male and two females) to Panna in 2009 (Table S3). The reintroduction program in Panna tiger reserve recorded better growth in the tiger population (40.8%) compared to that of Sariska tiger reserve (16% Table S3) perhaps due to more inviolate space available and less human disturbance in Panna compared to Sariska tiger reserve. Tiger supplementation was attempted in Satkosia tiger reserve in 2018 but it failed due to community opposition and poor status of tiger prey. Tigers have been successfully reintroduced in Nauradehi wildlife sanctuary and translocated to Satpura tiger reserve so as to reinforce low tiger density areas by the Madhya Pradesh Forest Department.
Tiger reintroductions/reinforcements attempted in India were done using (a) wild tigers captured and introduced into the wild at another site, (b) hand reared tigers introduced into the wild and (c) orphaned or abandoned cubs cared for in-situ by food supplementation and veterinary care. The survival beyond first year of wild tigers (n = 27; survival 72%) did not differ from that of hand reared tigers (n = 13; survival 77%) introduced to the wild. All cubs (n = 7) assisted in-situ by supplementation of food survived beyond the first year. Successful reproduction in the wild after introduction was recorded in 50% (n = 12), 55% (n = 9) and 75% (n = 4) of wild, hand reared and supplemented tigresses respectively. The success of introductions was not dependent on the origin (hand reared vs. wild caught) of the tiger being introduced, but more on the conditions at the site of the introduction. Introductions were more successful in areas with no or low tiger density, good prey density, good law enforcement mechanisms and friendly neighbouring communities. Introduction of tigers should be done only after proper assessment of these factors and only after these conditions are found to be optimal. Considering a kill rate of one medium-sized ungulate (50-80 kg) every 3-4 days by an adult tiger and realized finite growth rate (λ) of ~1.3 for ungulates (Duncan et al., 2007), as a thumb rule 450 ungulates to one adult tiger was considered as a minimum prey requirement (see Fuller, 1989). For reintroduction to be considered at a site, sufficient prey for at least 12-15 tigers was considered a prerequisite at the site (Table S3). With good protection and management, depressed prey populations at such sites would increase further along with the introduced tigers. Once reintroduced tigers start to breed, reintroduction programs often do not follow-up with continued reinforcement of additional tigers as per the reintroduction plan, resulting in poor genetic diversity in these newly established populations. Immigration, either natural through corridors, or through managed reinforcement is essential for long-term survival of these tiger populations.

| Assessment and monitoring
The four-yearly monitoring of tiger status based on modern animal abundance estimation science commenced since 2006, and covers all potential tiger bearing forests of India (~381,000 km 2 ). The monitoring protocol also evaluates the status of co-predators, prey and habitat. The protocol involves ground surveys by ~44,000 personnel who now collect information using mobile phone GPS enabled application MSTrIPES (Ecological module) in an occupancy (MacKenzie et al., 2017) and distance sampling framework (Buckland et al., 2005).
Spatial sampling units consist of forest beats (~15 km 2 ) or 25 km 2 grids (Jhala et al., 2020). Spatially explicit data are digitally recorded with photo-evidence of wildlife signs stamped with date, time and geographic coordinates, that are sampled as occupancy surveys; ungulate sightings on line transects; vegetation, pellet density and human disturbance are sampled on plots. Subsequently, camera traps are used to obtain data on photo-captures that are analysed in a spatially explicit capture-recapture framework (Borchers & Efford, 2008) to estimate densities of tigers and leopards. While for other species relative abundance is indexed (RAI) using number of independent photo-events corrected for sampling effort (trap-nights). All potential tiger habitats (tiger reserves, protected areas, reserve forests, Considering only the consistently assessed areas for all four population estimation cycles, the tiger population across India has been growing at a rate of 6 (SE 0.6) % per annum ( Figure S1). Besides the four-yearly national assessment, all tiger reserves (harbouring ~65%  (Table S1). Incidentally, the largest Asiatic elephant Elephas maximus populations too are found in these two habitats, highlighting the umbrella role of tiger conservation.
The strategy of prioritizing conservation of tiger source populations within each landscape and simultaneously connecting these sources amongst themselves and sink habitats through corridors, has resulted in tiger occupancy across suitable habitats within landscapes and helped recover these populations (Bisht et al., 2019;Walston et al., 2010).
Since substantial investment goes into tiger reserves, these reserves were evaluated for their management effectiveness every four years since 2006, by a team of independent evaluators. Tiger reserves are evaluated for 34 criteria covering the themes of status, appropriateness, resources, efficiency of effectiveness, outputs and outcomes following Hockings et al. (2006). Tiger reserves management effectiveness performance ( Figure S2, and data sources) was then Often Protected areas are considered as lacking sufficient economic justification by politicians, legislators and bureaucrats especially in countries like India where there is a high demand for land due to dense human population and poverty. In such cases, it becomes difficult to justify setting aside large areas as tiger reserves where usual extractive human use is prohibited. However, when tiger reserves are valued in terms of their economic contribution and are shown to be competitive in terms of monetary gains compared with other economic activities, it becomes easier to convince the decision makers of their importance. Select tiger reserves were evaluated for their tangible economic value based on standing crop, carbon sequestering, employment generation, direct profits and ecosystem services, amongst other values that could be monetized (Costanza et al., 2014). Based on these evaluations the value of annual flow of benefits from a tiger reserve ranged between 76,900 and 292,300 US$/km 2 (Verma et al., 2017). The language of economics appeals to decision makers and if appropriate safe-guards and caveats (to ensure that monetary value is not the sole criteria for evaluating Protected Areas) are appropriately communicated, support for tiger reserves is garnered across most sections of the society.

| WAY AHE AD
Within India, tiger conservation efforts and the associated increase in tiger populations is not evenly distributed across tiger habitats.
In many tiger habitats such as the North Eastern Hills bordering Myanmar, and in the states of Odisha, Jharkhand and Chhattisgarh, forest communities still consume bushmeat. In these areas, forests are almost devoid of wild prey and therefore of tigers. Wild ungulates are often poached here with the use of snares that do not discriminate between species, often killing large carnivores like tigers as well. Some tiger habitats in these same states are strongholds of left-wing extremist element (Naxalites) making management activities difficult to implement. Once habitats within these States are restored to law and order, and communities made economically secure they may be weaned off bushmeat. Prey and subsequently tigers would respond and recover since the habitat is good. Some biodiversity-rich and promising areas like Guru Ghasidas National Park, in this landscape, would benefit from focused conservation investment and should be notified as a Tiger Reserve. This would allow resources of Project Tiger to be accessed for incentivized voluntary relocation, community upliftment activities, prey and predator population reinforcement, amongst other activities that will hasten ecosystem recovery. Once prey populations are restored, another 1,000-1,500 tigers can be accommodated in these regions.
The establishment of Tiger Conservation Foundations, sharing of tiger-profits with buffer zone communities, and a handsome package for incentivized voluntary relocation have gone a long way in winning over local communities. Yet, more needs to be done for the communities that actually bear the costs associated with tiger conservation, especially when tiger populations expand beyond protected areas.
Tiger and associated eco-tourism have a lot to offer in this regard.
Besides a few jobs, profits rarely percolate to local communities from the tourist industry. Community-based home stays with a menu of options like trail walks, birding, night safari and campfire-based activities in the buffer zone and outside tiger reserves (which are prohibited in the core zone of tiger reserves) hold promise of attracting a different clientele of wildlife tourists, those who seek a greater immersion in the experience of the wilderness. Communities were observed to be more tolerant to carnivore damage and even encouraged large carnivores in their neighbourhood if they could realize remunerative livelihoods from them . Village consortiums that manage their lands for wildlife values and have rights to earn revenues from wildlife resources on their lands would be a major conservation paradigm shift for India . Some states like that of Rajasthan, have yet to enact legislation that trans- Pseudomelenistic tiger from Simlipal tiger reserve, (d) Golden tiger from Kaziranga tiger reserve, (e) White tiger in captivity whose ancestral origin was from the Rewa forests near Sanjay tiger reserve, (f) Light coat variant (almost white) from Western Ghats habitats, and predating different prey. There is sufficient evidence to suggest genetic differences amongst them due to vicariant events and local selection (Luo et al., 2019). For retaining the evolutionary potential of tigers as a species, we should strive to conserve the entire extant genepool, range of habitats, behaviours that are culturally inherited, and manage populations so as to retain their unique evolutionary trajectories while mitigating fragmentation caused by human activities. Based on genetic diversity, divergence and vulnerability to extinction, tiger populations have been prioritized for conservation within India (Kolipakam et al., 2019). Tiger populations of the North Eastern Hills and those of Southern Western Ghats were considered a conservation priority based on these criteria (figure 6 in Kolipakam et al., 2019). The NTCA has a standard operating procedure for guiding tiger reserve managers on sourcing tigers for reintroductions and supplementation (https://ntca.gov.in/docum ents/#sop1) so as to retain local adaptations while attempting to prevent inbreeding. A global conservation priority analysis of this kind is likely to prioritize tiger subspecies P. t. crobetti and P. t. jacksoni since populations of these two subspecies are genetically distinct and precariously low.
India has managed to increase its adult tiger population from around 1,700 in 2010 to about 3,000 by 2019 with tigers currently occupying close to 90,000 km 2 . This is no minor feat by global standards (Figure 1), since many tiger range countries that are economically better off than India, have recorded declines in their tiger populations and occupied range. Tiger conservation is no rocket science, the road towards recovery begins with the acknowledgement of problems and subsequent management on the scientific principles of conservation biology. Misinformation regarding tiger population numbers by the park authorities, that were subsequently defended by the State bureaucracy led to poaching-driven extinctions in Sariska and in Panna tiger reserve (Chundawat, 2018;Narain et al., 2005). India, Nepal and Bangladesh took a bold step and corrected their national tiger population estimates based on science in 2008, 2011 and 2015 respectively (Table S1). Many range countries still rely on erroneous or misinformed tiger estimates that are obtained through unreliable methods (Table S1). Often the inertia to acknowledge a problem by various levels of bureaucracy makes efficient and timely conservation interventions difficult.
Major credit for India's tiger recovery can be attributed to a multidisciplinary approach fostered by trust and team spirit between wildlife managers, bureaucrats, activists, scientists and legislators.
However, conservation practitioners need to put in more effort so as to mainstream biodiversity concerns with the agenda of other government agencies like mining, roadways, electricity, irrigation, public works development agency, etc. Often the objectives of these agencies clash with those of conserving tigers and their ecosystems. It is absolutely essential to engage with and sensitize these public sectors regularly to conservation needs, so that one arm of the government does not undo what the other does, using the same public funds.
The reforms mentioned above would not have been possible without the political will at the highest level of Governance. This commitment to conservation of India's national animal as a flagship and umbrella for the ecosystems it inhabits, transcends political party lines. In the previous decade, Prime Ministers Manmohan Singh and Narendra Modi have shown their commitment towards tiger conservation and pledged resources to make it a reality on the ground. Range countries need to ensure that tiger (and biodiversity) conservation becomes a priority national agenda with appropriate funds allocated by Governments. Funding from national and international conservation agencies/NGO's, although important, should target specific weak links where Governments find it difficult to use public funds. In the post COVID pandemic scenario, a depressed global economy will put huge pressures on tiger habitats, some of which, are rich with coal and mineral deposits. International funding support, for this conservation-dependent species becomes even more important in this period of crisis, when national funding priorities of most range countries will likely shift towards exploitative development activities. How India and other tiger range countries manage to balance their economic growth without compromising the conservation of their natural heritage and ecological security, will decide the fate of wild tigers.
Currently, tigers across their range are primarily threatened by direct poaching and secondly by prey depletion (Chapron et al., 2008).
As long as there is a demand for tiger parts and products, there will be poachers who will be willing to take high risks. The global community needs to build pressure to eradicate the demand for wildlife parts and products, especially in light of the current pandemic that has likely arisen due to this trade that not only threatens endangered species but even puts human survival at risk.

ACK N OWLED G EM ENTS
We thank the forest department field staff whose hard work and dedication has resulted in the recovery of wild tigers. Swati Saini is thanked for map preparation. Lex Hiby and Rajan Amin are acknowledged for their assistance with ExtractCompare and MSTrIPES. provided technical and scientific support for tiger conservation policy and monitoring. All authors read and commented on the MS.

DATA AVA I L A B I L I T Y S TAT E M E N T
No primary data were generated for this study. All data sources are appropriately cited in the paper.