Jaguar i naturen

Understanding hybrid animals better



In the series of two hybrid blogs, we examine how changing and evolving ideas on hybrid animals can help us understand them better.

Hybrids are the future of the world

The recent Netflix series, Sweet Tooth, has a memorable sequence showing a child named Bear saying, “Hybrids are the future of the world, therefore we as children, have taken a vow to protect all the hybrids.” In this feature, in two parts, we examine how changing and evolving ideas on hybrid animals can help us understand hybrid animals better, including better perspectives on how hybrid animals are represented in wildlife trade.

The popular television series, that depicts hybrids between humans and animals, has given rise to renewed debate on the nature of hybridisation as a phenomenon and if, in the future, there will be a revised understanding of hybrid organisms, both plant and animal.

In July 2016, scouring through the archives of the National Library in Kolkata, India, an information scientist and a librarian laid their hands upon a rare photograph published in 1980 in the daily newspaper, The Statesman.

The photograph was that of a male litigon. It was described in an accompanying news report as a hybrid of a male Asiatic lion (Panthera leo persica) and a female tigon (hybrid of a male tiger Panthera tigris and a female African lion Panthera leo of unknown subspecies) from the Alipore Zoological Gardens in Calcutta (now Kolkata).

The litigon was named Cubanacan by Jose Lopez Sanchez, the erstwhile Cuban Ambassador to India and photographed on the cub’s first day of public viewing in the zoo.

Cubanacan, the progeny of a lion and a tigon, was born at the Alipore Zoo in Kolkata, India, on March 7, 1979 and was the only surviving cub of his litter of three.


A photo of the litigon Cubanacan in Alipore Zoo published in the March 12, 1980 edition of The Statesman, Calcutta. Credit Subhodip Bid and Payel Biswas

A photo of the litigon Cubanacan in Alipore Zoo published in the March 12, 1980 edition of The Statesman, Calcutta. Credit Subhodip Bid and Payel Biswas


Mon Pobon magazine editor Rahul Majumdar recovered three more photographs of Cubanacan, the litigon, in Alipore Zoo on August 8, 2018. This added to the invaluable archive of the much-maligned and neglected phenomenon of cross-breeding big cats in Alipore Zoo in Kolkata.

Alipore Zoo had embarked on a 15-year endeavour to hybridise lions and tigers, an effort that created Cubanacan’s tigon mother, Rudrani, and her sister, Rangini, several years earlier.

Cubanacan was born after 15 years of hybridisation attempts that started in 1964 at the Alipore Zoo. The zoo reportedly produced its first hybrid big cat, a tigon called Rudrani, on October 13, 1972 in the sixth litter of a female African lion Munni and a male tiger Munna.

A second tigon, Rangini, was born on March 8, 1974. Both tigons were named by the then chief minister of West Bengal, Siddhartha Shankar Ray. A pioneering scientific success for India and even the rest of the world, Cubanacan was widely regarded as the first litigon born in the world.

Cubanacan’s remarkable genetic makeup sparked interest and enthusiasm in India and around the globe. The fascination with hybrid cats continued as Rudrani produced four more litigons in subsequent years.


A photo of the litigon Cubanacan of Alipore Zoo in the 1985 Guinness Book of Records. Photo: Alipore Zoo and Karl Shuker

A photo of the litigon Cubanacan of Alipore Zoo in the 1985 Guinness Book of Records. Photo: Alipore Zoo and Karl Shuker

There is now evidence that these experiments were led by a scientific quest to determine if hybrids could be fertile, a question that struck at the heart of the notion of biological species.

At the time, the very definition of species hinged on reproductive isolation. Though probing at a research question, concerns surfaced about artificially creating animals not found in the wild as freaks for public curiosity.

There were also claims of animal cruelty during the process, an allegation that has come to the forefront in the current effort to ban cross-breeding of big cats in the United States.

In the midst of this controversy, hybrids still command ample public attention. The 2017 Guinness World Records (formerly the Guinness Book of Records) ranked, Hercules, a liger [lion x tigress] at the Myrtle Beach Safari in South Carolina, US, the world’s largest big cat.

Cubanacan was also once the world’s largest big cat, who, according to Guinness in 1985, weighed 363 kg (800 pounds), stood 1.32 metres (52 inches) at the shoulder and measured 3.5 m (138 inches) in length.

Given the aversion to hybridisation and the subsequent 1985 ban on cross-breeding big cats in India, it appears that Cubanacan’s memory was purposely forgotten.

The scientific aspects of the big cat hybridisation 

Big-cat researcher, Ratan Lal Brahmachary was deeply interested in studying the scientific aspects of the big cat hybridisation experiments carried out in Alipore Zoo. In a personal letter written to me (Shubhobroto Ghosh) on August 28, 2015, Professor Brahmachary expressed his views on Cubanacan the Litigon and other hybrid big cats:

Received your numerous queries on Cubanacan. I think the large size is a result of hybrid vigour. I saw ligers in Khartoum Zoo, they were also large. White tigers are also larger than the normal-coloured ones. Unfortunately, no scientific study on Cubanacan was carried out. I was most interested to know whether his urine / marking fluid was aromatic like that of the tiger or like the lion. No comparison of blood proteins or anything of that sort was attempted. Regarding research, the zoo had a dog in the manger policy as I learnt in 1980s.

In his book, Bagh O Tar Gyati Goshthi, published in 2011, Brahmachary wrote about a triple-hybrid, a lion X leopard X jaguar hybrid. Recently, zoologist and author and Guinness World of Records consultant, Karl Shuker, retrieved a long lost photograph of this hybrid creature at London Zoo on  May 22, 2017.

Karl Shuker recalled the eventful history of the truly remarkable big cat — viz, an adult female three-way hybrid, aptly dubbed Uneeka. Her parents were a male lion and a female jaguar x leopard hybrid, thereby making her a lijagupard.

She was originally claimed to be a new species and exhibited in London Zoo for just over a fortnight during April 1908. She was then purchased by Glasgow-based circus / menagerie-owner Edward H Bostock on May 2 for the very sizeable sum of 1,030 guineas and taken by him to Glasgow. Uneeka was allegedly killed just over a year later by a lion that supposedly broke through from its own cage into Uneeka's.

Karl Shuker was both delighted but also very startled recently. That is because he discovered a black-and-white photograph of Uneeka in a vintage issue of a London weekly magazine entitled the Illustrated London News.

The image showed her sitting inside a cage. The zoologically-invaluable photo was one of a series of photographs depicting various London-themed subjects that had lately been in the news, all presented together on a single page in the May 9, 1908 issue of this magazine.

Like Cubanacan, Uneeka’s very existence shows that fertile hybrids very much exist and existed during the time when it was popularly presumed that all hybrids were sterile like the mule (Records of fertile mules exist, by the way).

 Uneeka, the lion-leopard jaguar hybrid as photographed on May 9, 1908. Photo:  Illustrated London News, discovered by Karl Shuker

Uneeka, the lion-leopard jaguar hybrid as photographed on May 9, 1908. Photo:  Illustrated London News, discovered by Karl Shuker


The hybridisation debate in biology is important. Researchers at Texas A & M University in the US and the Pontifícia Universidade Católica do Rio Grande do Sul in Brazil have come a step closer to understanding the rich evolutionary history of the cat family.

In a paper, featured on the cover of Genome Research journal in January 2016, the researchers constructed extensive family trees of the 38 cat species, which illustrated maternal, paternal and biparental lineages within the cat family.

However, they found that lineages are not completely linear. Instead, this study revealed that feline ancestry has been shaped throughout its evolutionary history by hybridisation.

The recent finding of the litigon photograph warrants a fresh look into the biological significance and ethical connotations of the classic experiments performed at the Alipore Zoo.

This assumes importance following recent suggestions that interspecies hybridisation may be a natural and significant biological phenomenon. Critical questions have also been raised on the validity of the widely accepted biological species concept that has traditionally defined ‘species’ as reproductively isolated taxa that cannot mate with other taxa to produce viable offspring.

Hybridisation -- threat to endangered taxa?

Conservationists suggest that hybridisation between different species and subspecies is among the many threats to endangered taxa, alongside habitat destruction and degradation, poaching, pollution and illegal wildlife trade.

This view has been endorsed by several international organisations, including the International Union for the Conservation of Nature (IUCN), the World Association of Zoos and Aquariums (WAZA) and many national agencies.

It has led to widespread condemnation of hybridisation in captivity and in the wild. The extreme abhorrence for hybridisation has also reportedly resulted in the euthanasia of hybrid animals in zoos.

Today, research by evolutionary biologists like James Mallet of Harvard University shows that on an average, at least 10 percent of animal species and maybe 25 per cent of plant species are known to hybridise in nature.

The fraction of species that hybridise may, however, be much higher in rapidly radiating groups. A fundamental concern is the widespread view which considers inter-species breeding as a form of ‘genetic pollution’.

This view fuels the widespread academic notion that species are reproductively isolated units and that all events of hybridisation are undesirable accidents, with hybrids (such as sterile mules) being evolutionary dead-ends.

This concept has become a part of standard discourse on biological hybrids. Do these positions, both in the biological sciences as well as in conservation, actually stand up to scrutiny?

The definition of the biological ‘species’, as a fundamental category of biological organisation, has been a long-standing conundrum in biology.

The most widely accepted definition, proposed by Ernst Mayr, suggests that species, at least in a taxonomic sense, are groups of interbreeding natural populations reproductively isolated from other such groups.

This definition also explains why members of a species resemble one another closely and differ from other species in biological traits.

The link between species and gene pools or metapopulations

A number of definitions arose during the period of the Modern Evolutionary Synthesis, including Mayr’s. But, the most comprehensive and well-subscribed concept emphasised the link between species and gene pools or metapopulations.

These concepts, however, differed in certain properties deemed necessary for an independently evolving metapopulation lineage to be considered a separate species.

These characteristics included specific mate recognition systems, ecological distinctiveness, monophyly, formation of distinctive phenetic classes and, importantly, intrinsic reproductive isolation.

These alternative definitions of species demanded that one or more of these characteristics be considered necessary properties of species and this, in turn, led to debates on which metapopulations could be considered as distinct species.

A historical analysis of speciation processes shows that these so-called properties of species may have originated at different times and followed different developmental paths of evolution.

For instance, in the case of reproductive isolation, several pre-and post-zygotic mechanisms (genetic, chromosomal, morphological, anatomical or behavioural) may be responsible for species recognition and/or breeding incompatibility between members of different metapopulation lineages.

Importantly, these mechanisms could have arisen by chance — through mutations or chromosomal rearrangements and/or facilitated by migration or genetic drift.

They may have also evolved out of necessity, by natural selection, under conditions such as sympatric speciation, the process through which new species evolve from a single ancestral species inhabiting the same geographic region.

Moreover, the evolution of these mechanisms may not be related to other evolving properties of a potential species, such as shared ecological niches, monophyly or fixed qualitative or quantitative traits that distinguish a metapopulation lineage from another as distinct species.

Thus, two metapopulation lineages may have differentiated into two distinct species by any of these criteria and yet not have evolved intrinsic reproductive isolation. And this may have happened in the course of the evolutionary histories of the tiger and the lion.

Why has this first-hand evidence of the breeding viability between different kinds of hybrids failed to attract the attention of biologists? Why have these remarkable biological events not been acknowledged with the scientific importance they deserved?

The answers to these questions may constitute classic examples of the way science often progresses and reflect the sociology of scientific discovery and experimentation in particular regimes of dominant scientific ideologies.

There is need for an inquiry into the motivation and the decision-making process that led to the creation of the tigon and the litigon in the Alipore Zoo between 1964 and 1979.

Was it plain curiosity or was it a scientific endeavour to better understand mammalian genetics and evolutionary forces at work?

An official statement of the time states:

Meanwhile, attempts are afoot to induce companionship between Rudrani and an Indian lion called Devavrata. Signs of mutual acceptability are discernible and should compatibility result perchance, birth of a triple hybrid may not be that improbable as believed so far.

Amarendra Nath Guha, then director of the Alipore Zoo, was also quoted in a newspaper as saying, “(Cubanacan was) an ambitious hybrid of a type believed to be improbable so far”


Apart from the big cat hybrids bred in Alipore Zoo in Calcutta, there is also the case of Ranji. 

He was a huge male lion x tiger hybrid who was bred by Prince Ranjitsinhji, Maharajah Jam Sahib of Nawanagar in Gujarat and presented by him to the Zoological Gardens in London in Regent's Park in 1924. 

The Illustrated Sporting and Dramatic News of London, November 19, 1927, carried a long article on the London Zoo tigon entitled, ‘FRIENDS IN CAPTIVITY-- VII. The Tigon. By S. L. Bensusan. Illustrations by L.R. Brightwell.’

Published by Sarah Hartwell on her website on hybrid animals, the entry states that, “This may not be the only experiment of the kind that has been carried out by the ruler through whose menagerie the tigon has reached London.”

One of the authors (Shubhobroto Ghosh) has recently been able to verify the nature of this experiment with His Highness Shri Jamsaheb Shatrusalyasinhji of Jamnagar in Gujarat.

Ranji was the product of an experiment to test the reproductive compatibility of big cats like lions and tigers and similar interests were harboured by London Zoo more than one hundred years ago. 

These endeavours now carry added scientific value in the aftermath of papers revealing the hybrid origin of big cats in the wild such as the study done by William Murphy of Texas A & M University in USA in 2016.


Ranji, the lion-tiger hybrid bred by Jam Sahib, the Maharajah of Nawanagar and presented to London Zoo. Photo: Illustrated London News 1924 discovered by Sarah Hartwell


 Ranji, the lion-tiger hybrid bred by Jam Sahib, the Maharajah of Nawanagar and presented to London Zoo. Photo: Illustrated London News 1924 discovered by Sarah Hartwell

Hybridisation and value-driven science

There is now extensive evidence on other hybridisation cases between several distantly related cat species in captivity.

Many common domestic cat breeds, such as the Bengal — a hybrid between the domestic cat and the Asian leopard cat — are also believed to be of hybrid origin.

In fact, feline ancestry appears to have been shaped by natural hybridisation events throughout its evolutionary history.

Increasingly, evidence is being uncovered to show how profoundly hybridisation has influenced the natural evolution of taxa as diverse as butterflies, sharks, finches, parrots, dolphins, bears, wolves, old world primates or even modern humans.

For example, research by evolutionary biologist James Mallet shows that hybridisation between species of Heliconius butterflies seems to be a natural phenomenon. There is no evidence that it has been enhanced by recent disturbance of habitat by humans.

Molecular clock dating suggests that gene exchange by way of hybridisation may continue for more than three million years after speciation. In addition, one species, Heliconius heurippa, appears to have formed as a result of hybrid speciation. Introgression may often contribute to adaptive evolution as well as sometimes to speciation itself, via processes like hybrid speciation.

Geographic races and species that coexist in sympatry therefore form part of a continuum in terms of hybridisation rates.This finding agrees with the view that processes leading to speciation are continuous, rather than sudden and that they are the same as those operating within species, rather than requiring special punctuated effects or complete allopatry.

Although not qualitatively distinct from geographic races, nor ‘real’ in terms of phylogenetic species concepts or the biological species concept, hybridising species of Heliconius are stably distinct in sympatry and remain useful groups for predicting morphological, ecological, behavioural and genetic characteristics.

Interbreeding between animal species can lead to offspring less vigorous than either parent — if they survive at all. But the combination of wolf, coyote and dog DNA that resulted from a reproductive necessity generated an exception.

The consequence has been booming numbers of an extraordinarily fit new animal spreading through the eastern part of North America. Some call this creature the eastern coyote.

Others, though, have dubbed it the ‘coywolf’. Whatever name it goes by, Roland Kays of North Carolina State University, in Raleigh, reckons it now numbers in the millions.

Whether the coywolf actually has evolved into a distinct species is debated. Jonathan Way, who works in Massachusetts for the National Park Service, claims in a paper that it has.He thinks its morphological and genetic divergence from its ancestors is sufficient to qualify. But many disagree.

One common definition of a species is a population that will not interbreed with outsiders. Since coywolves continue to mate with dogs and wolves, the argument goes, they are therefore not a species.

But, given the way coywolves came into existence, that definition would mean wolves and coyotes should not be considered different species either —and that does not even begin to address whether domestic dogs are a species, or just an aberrant form of wolf.

In reality, ‘species’ is a concept invented by human beings. And, as this argument shows, that concept is not clear-cut.

What the example of the coywolf does demonstrate, though, is that evolution is not the simple process of one species branching into many that the textbooks might have you believe.

Indeed, recent genetic research has discovered that even Homo sapiens is partly a product of hybridisation. Modern Europeans carry Neanderthal genes and modern East Asians the genes of a newly recognised type of early man called the Denisovans.

Exactly how this happened is unclear. But maybe, as with the wolves of southern Ontario, it was the only way that some of the early settlers of those areas could get a date.


Acknowledgements :  Rahul Majumdar, Dr Anindya Sinha, senior scientist, National Institute of Advanced Studies, Bangalore, Piyali Chattopadhyay Sinha, Ashish Samanta and Shibaji Bhattacharya, Alipore Zoological Gardens, Kolkata, Payel Biswas of Institute of Urban Transport, New Delhi, Subhodip Bid of National Library, Kolkata and Anirban Chaudhuri all contributed in locating the historical records of hybrid big cats in Alipore Zoo in Kolkata. The authors extend their gratitude to Dr M K Ranjitsinh and Ms Ekta Sodha, CEO of Cadmus Sodha Schools in Jamnagar for their help in confirming the record of Ranji, the tiger X lion hybrid in the zoo of His Highness, the Jamsahib Ranjitsinhji, Maharajah of Nawanagar, Gujarat, India.



Shubhobroto Ghosh is author of the Indian Zoo Inquiry and the book, ‘Dreaming In Calcutta And Channel Islands.’ He is now Project Manager of Wildlife at World Animal Protection in India


Karin Saks is a primatologist based in South Africa. Since 1997, Karin has been involved in the fostering and rehabilitation of orphan baboons, the caring of injured monkeys who have been returned to the wild and has worked towards a harmonious co-existence between these primates and humans. She has many publications on primates and has been the subject of television programmes and films