PROLOGU E Captain Thomas Williamson, the author of the epic book about India’s animals, Oriental Field Sports, says that in the 1780s, while pig-sticking, one was likely to encounter tigers that had strayed into the open, but never lions, or for that matter, cheetahs. He went on to state that, after spending the last two decades of the eighteenth century roaming the wilderness of India, he believed that there were no lions in Hindustan. For me this was an excellent clue regarding the state of the lion during that time, which was, then, a much debated topic. In his observations of the same period, Thomas Pennant says of lions: ‘Those who deny that those animals were natives of India, assert that here was a royal menagerie and that the breed was propagated from the beasts which had escaped.’ The debate about the origins and prevalence of lions and cheetahs in India must have been vigorous in the eighteenth century and later but I was intrigued by the fact that it hasn’t been talked about much in recent years. In fact, in the twentieth century, most serious observers just took it for granted that lions and cheetahs were indigenous to India. In his book, Wild Animals in Central India, published in 1931, A. A. Dunbar Brander writes that there were no lions in Central India and goes on to say of the cheetah that, ‘The Hunting Leopard (Cynalurus jubatus) has now almost entirely disappeared from the province without apparent reason, and I only know of three animals having been procured in the last twenty years.’ As we will see in the course of this book, it is likely that what Brander referred to must have been three trained cheetahs imported from Africa. Why is it that the comments about lions and cheetahs and their rarity remain unchanged over hundreds of years, especially from people who travelled across India in search of game? Why are pictures of lions as hunting trophies so rare and the few that exist tend to have been taken 13
after 1886 in Gir? Where are the cheetahs in the hunters’ bags? After all, according to commentators, hunters were responsible for destroying both these species. As I researched my thesis, I grew convinced that I had stumbled upon the biggest myth perpetrated about these two species in India. Tigers and leopards were everywhere in historical records, whether written, photographed or depicted in other visual forms, but lions and cheetahs were almost invisible. Even on those rare occasions when lions and cheetahs were seen, their behaviour appeared forced and unnatural compared to what I had witnessed of both species in Africa. As I researched into the prehistoric era (eight to ten thousand years ago) through extraordinary books like Erwin Neumayer’s Lines on Stone: The Prehistoric Rock Art of India—which reveals the absence of ‘maned’ felines on rock—as well as through conversations with my aunt, Romila Thapar, and followed the thread of my argument through the pre-Christian, pre-Islamic, Mughal and colonial periods, my conclusion gradually began to coalesce. In the course of the nearly four decades that I have studied the Indian tiger, I have had occasion to add to the sum of knowledge on that magnificent predator, but this serendipitous book on the Indian lion and cheetah has been stimulating in new and exciting ways. Is there an evolutionary or genetic basis to my thesis that goes beyond the historical record? The tiger, lion, leopard and jaguar are the four big cats that belong to the genus, Panthera. The reason they are grouped together is because of their ability to roar, even though tigers do not roar like lions and leopards. This ability is made possible by the vibration of the thickened vocal folds below the vocal chords in the larynx. This group of cats has probably descended from mammals called Miacids (Miacidae), the first true carnivores, which crept around treetops nearly 50 million years ago; at this time they were thought to be the size of martens. About 20 million years ago, a group of mammals called the 14
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pseudaelurines evolved—these are, in all likelihood, the direct ancestors of all wild cats today. It is believed that two million years ago, the common ancestor to the Panthera cats looked like the leopard does today. After a series of genetic comparisons, zoologists believe that the tiger was the first to diverge from the common Panthera ancestor, followed by the lion. Because of this fact, lions, leopards and jaguars have more in common with each other than with the tiger. Tigers and lions look very similar when they are stripped of their stripes and this makes it very difficult to prove which one came into existence when. Hence, it has been very difficult for experts to trace the evolutionary path of tigers and lions. When it comes to separating a species of cat like the tiger or the lion into a subspecies, things that are taken into account are range or physical characteristics. Biologist Andrew Kitchener states: ‘Sometimes a species’ morphology changes gradually over its geographical distribution so that at the extreme of their range animals may look completely different. This is known as a cline, and even though individuals on the opposite end of the cline may look very different, it is inappropriate to regard them as belonging to different subspecies. There is always gene flow throughout the geographic range, albeit tempered by natural selection for particular morphotypes or perhaps reflecting a pattern of broad hybridization between two populations that differentiated previously in isolation. Sometimes local populations are separated from one another by geographical barriers, such as a river or a mountain range, and they differ in their appearance such that at least 75 per cent of one population can be distinguished from 100 per cent of the other. In such a case it is possible to recognize two distinct subspecies or geographical races which can be given different scientific names. Finally, in many cases, local populations of morphologically different animals, may meet but have a narrow hybrid zone between the two, where animals of mixed appearance occur. Therefore there is always some limited gene flow between most subspecies. Although each animal species and subspecies has its own scientific name, science was often not involved in determining their distinctiveness. Many scientific names
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date back to between the late eighteenth and early twentieth centuries and are often based on a single or a few specimens.’ Traditionally, twelve subspecies of lions were recognized based on their geographical distribution, size, shape, and so on (these divisions were based on an analysis of zoo specimens). In the very recent past, eight lion subspecies have been more or less accepted although even within these some are considered invalid. These subspecies are: the Asiatic lion (Panthera leo persica), once widespread across Turkey, southwest Asia and India; the Barbary lion (Panthera leo leo), originally found from Morocco to Egypt and now extinct; the West African lion (Panthera leo senegalensis); the Northeast Congo lion (Panthera leo azandica); the East African or Masai lion (Panthera leo nubica); the Southwest African lion (Panthera leo bleyenberghi); the Southeast African or Transvaal lion (Panthera leo krugeri); and the Cape lion (Panthera leo melanochaita), now extinct. New research suggests that the modern lion can be divided into three geographical populations: North African and Asian, Southern African and Middle African. I am sure that in the future this will change again but as far as my historical research is concerned, the Indian or Gir lion is a mixture of some of the above groups.  The cheetah probably evolved in Africa, but some 10,000-12,000 years ago it faced a severe genetic bottleneck in its population resulting in the near extinction of the species; although it survived, a high incidence of inbreeding took place within the remaining population. Till recently, many believed there were at least five subspecies of the cheetah. These are: the Asiatic cheetah (Acinonyx jubatus venaticus) that roamed across Iraq, Iran, Israel, Jordan, Oman, Saudi Arabia, Syria, Russia, India and many other countries in the region; the Northwest African cheetah (Acinonyx jubatus hecki); the Eastern African cheetah (Acinonyx jubatus raineyii); the Southern African cheetah (Acinonyx jubatus jubatus) and the Central African cheetah (Acinonyx jubatus soemmeringii). The issue of the subspecies of the cheetah is still unresolved as many believe the North African and Asiatic cheetahs are the 16
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same but others hold that the Asiatic and African are different. The confusion arises from the fact that genetic variations among cheetahs are very slight and I am sure, with time, this classification will undergo further change. Be that as it may, I believe the cheetah in India has been imported from other countries. Given the constantly shifting classification of lions and cheetahs based on the evolutionary record, is genetics an effective determinant of whether a subspecies is distinctive or not, especially with regard to the Indian lion and cheetah? Unfortunately, genetic studies on these animals thus far have proved inconclusive and much more work needs to be done to find the answer. What I believe—and this is what my co-authors and I explore throughout the book—is that lions from Persia and Africa were being imported into the country 2,500 years ago (and then on) to meet the demand of Indian royals, and being bred and propagated as court symbols and for hunting; this imported animal was erroneously called the Asiatic lion. The story of the cheetah is much the same but its inability to breed in captivity meant that many more had to be imported. Both species in India were genetic mixtures of animals brought in from elsewhere, and their own inbreeding; their genetic makeup can be best described as a khichdi of genes. This premise of mine requires new research and DNA analysis of both ‘subspecies’. As I am not a geneticist, let me quote Stephen J. O’Brien who, in his eye-opening book, Tears of the Cheetah: The Genetic Secrets of Our Animal Ancestors says, ‘[The] physical traits in Asian lions are manifestations of extremely severe inbreeding in their very recent past. The evidence for our conclusions was encrypted in their genes.’ He writes that Asian lions from the zoo or Gir forest had ‘virtually zero genetic diversity’ compared to the African lion. ‘The incidence of malformed sperm was 66% in Gir lions on average, compared to 50% in crater lions and 25% in Serengeti lions. Gir lions had five times fewer motile sperm per ejaculate than Serengeti lions and a tenfold reduction in serum testosterone levels.’ The low level of testosterone not only increases the quantity of malformed sperm in a sample of semen, but also affects the development of the mane. ‘The Gir lion males
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had become “feminized” by their history of inbreeding.’ Stephen O’Brien was very clear that the DNA of the Gir lions was so similar that it was as if they were all clones or identical twins. After analyzing samples taken from Asian lions being bred in 38 American zoos, O’Brien stated: ‘When we considered the known mutation rate for microsatellites, we calculated that it would take around three thousand years for a severely bottlenecked population…to reconstitute as much new variation as we were seeing in modern Gir lions. The bottleneck that compromised the Gir lions’ genetic variation dated back not just one century but three millennia!’ Whatever might have happened around 2,500 to 3,000 years ago to cause severe inbreeding in the lion, this was also the time where I believe the lion as an exotic animal entered India and was propagated by man. O’Brien was also clear that the Asiatic lion being bred in the American zoos was a ‘mongrel’; instead of being a ‘pure strain’, it had two African forebears among the five original founders. As far as I was concerned, this was what connected the Asiatic lion to Africa. O’Brien believed that the Gir peninsula (which is the only part of the country where the Asiatic lion is found) was like an island 2,500 years ago and the isolated population of lions in it suffered from severe inbreeding. Many others have agreed that this area along with Kutch was once an island, as were parts of Kathiawar. Romila Thapar states: ‘Even if this was lion country, there seem not to have been lions here. If there were lions here, why do they not occur on Indus seals, since the area had many Indus cities and settlements, such as the city of Dholavira on the edge of the Great Rann of Kutch? It would seem that there were no lions in this area during the period of the Harappan cities, around the mid-second millennium BCE.’ If there were no lions 4,500 years ago and they remained rare in the periods that followed, be it the Islamic, Mughal or British periods, then it must have been an exotic import all along as there is no gradual decline in the species. I believe that the lion came to India just before or with Alexander’s invasion of India around the third or fourth century 18
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BCE. It was from this time onwards that lions were bred, either in designated hunting parks or royal zoos, just like the Gir lion was bred in the late nineteenth century. The breeding of lions in ancient times was for the hunt, or to keep in menageries or in the court of the king as symbols of royal power. A recent paper by Barnett, Yamaguchi and Cooper entitled ‘The Origin, Current Diversity and Future Conservation of the Modern Lion (Panthera leo)’ states: ‘Over its natural range the modern lion may be evolutionarily subdivided into three geographic populations: North African-Asian, southern African and middle African, determined by expansions and contractions of mid latitude deserts and low latitude forests. The former two are characterized by relatively simple mtDNA haplotype structure, and the latter by a diverse haplotype mosaic. The North African-Asian population is separated from the Sub-Saharan African populations by the vast Sahara Desert, and its skull is morphologically distinguishable from those of Sub-Saharan lions (Hemmer 1974), suggesting its distinct taxonomic status.’ The interesting aspect of this piece of research is that these scientists put the North African and Asian lion into a single subspecies. In other words, there was just no ‘separate’ Asiatic lion. László Bartosiewicz reiterates this fact by saying: ‘Modern lions are closely related. Craniological traits of the now extinct North African Barbary lion (Panthera leo leo linnaeus, 1758) and Asiatic lions (Panthera leo persica meyer, 1826) are very similar. There must have been a contiguous population inhabiting North Africa and Asia.’ Even though the genetics of the species is complicated and necessarily the preserve of scientists, my research into some of the key findings in this area only strengthened my suspicions that the lion in India was an exotic alien. Where the cheetah in India is concerned, we should track back to O’Brien and his basic premise, where he argues that the biggest bottleneck in the history of the species was 10,00012,000 years ago when there were major changes and upheavals across the planet. He writes that the cataclysm that claimed saber-tooths, mastodons and so on almost decimated the cheetah
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as well; it was that ‘narrow escape from extinction’ that left its genetic mark of uniformity on the survivors. If, as O’Brien postulates, the major population of the cheetah in Africa itself is narrowly genetically diverse because of a ‘population bottleneck’ the species passed through millennia ago, I think it is extremely unlikely that a genetically distinctive subspecies could have existed in the subcontinent. We will explore this at greater length in my chapter on cheetahs. I believe there was never an ‘Asiatic cheetah’. Rather, this animal was an imported royal pet that escaped into the wild on several occasions and may, at times, have created small feral populations. The evidence that my co-authors and I have unearthed in the course of our research and study has duly underlined the basic premise that I began this book with—that the Indian (or Asiatic) lion and the Indian (or Asiatic) cheetah are not distinctive subspecies but are exotic aliens that live (or lived) in the land of the tiger. For me, this fact was made amply clear as I ploughed through hundreds of books and research papers on both the animals. The numbers just didn’t add up—I found fewer and fewer encounters with lions and cheetahs the more I read. Descriptions of both species were rare, and both seemed to lack a ‘tradition’ in India. Tigers and leopards were all over the place; lions and cheetahs in comparison were barely visible in the pages of history. Till the end of the nineteenth century, there were more lions seen in royal menageries than in the wild and hundreds (if not thousands) of cheetahs on leashes were recorded as being readied for coursing on bullock carts, but not even a handful were encountered in the wild. This book is the story of our search and raises serious questions about the ancestry and place of the lion and the cheetah in their natural state in India. If for some reason the premise I have put forward is proved to be wrong, I will be happy to be corrected. The only thing that matters is that the provenance of these two amazing animals is conclusively proved. I hope that the book provokes even more research into these incredible animals and their history in India.
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Two Jesuit priests in black caps and dark robes were among the first Europeans to visit the Mughal court at the end of the sixteenth century CE.They are laden with gifts including two cheetahs that are possibly for the Mughal prince, Salim. I believe that, in the absence of wild populations, hundreds of cheetahs were brought to India as gifts and tributes from visiting dignitaries (1590-1600).
Top: Dionysus, the God of the grape harvest, astride a lion and handing out wine. Gods preferred lions to ride on and this mosaic is from the museum of El-Jem in Tunisia (Second century CE). Bottom: In the absence of lions, tigers were ridden, as seen here with the Nawab of Jhajjar astride a tiger. Many rulers probably believed that taming or overcoming lions and tigers would give them supernatural powers (Twentieth century).