Thursday, April 25, 2013

Headless fight for Gir lions.

Gujarat government tried to defend its stance on retaining its lions but its wildlife board, which was meant to advise it on the issue, stood unconstituted since December 2012; lapse detected at a meeting this week
Hitarth Pandya
Posted On Thursday, April 25, 2013 at 08:57:59 AM

A high-power committee meeting regarding the issue has been scheduled for April 25
Ahmedabad - The Gujarat government harped on retaining its ‘pride’ and preserving its ‘unique heritage’ while arguing against the shifting of Asiatic lions from Gir in Saurashtra to Kuno Palpur in Madhya Pradesh, but it forgot to reconstitute the State Board for Wildlife (SBWL), meant to advise it on the issue. It realised its lapse only after a meeting was called to discuss the lions’ translocation.

Quickly, three MLAs were nominated to the board, as required by the SBWL statute, and invited to attend the meeting re-scheduled for April 25. State Forest Minister Ganpat Vasava said, “It is an important meeting and Chief Minister Narendra Modi is the chairman. MLAs, members from NGOs and others are board members. I’m not sure who the three MLAs are.

I am at my village and I can find it out later on,” he said. Senior officials in the forest department pressed the panic button on Monday when they realised that three MLAs who were part of the SBWL were not eligible to take part in the crucial meeting to discuss legal issues pertaining to shifting of lions.

When decision was taken to hold the meeting on April 25, some senior members realised that as per the constitution of the SBWL, it is necessary to re-appoint the MLAs on the committee as the term of the old ones gets dissolved after the Assembly elections in December 2012.

The state never took the necessary steps after the elections even as the case was on in the Supreme Court. The letter sent to SBWL members reads, “This is to bring to your kind notice that a number of members have requisitioned an urgent meeting of the State Board for Wildlife for discussing the directions issued by the Hon Supreme Court of India in judgment dated April 15, 2013, allowing the translocation of Asiatic lions from Gujarat to Madhya Pradesh.

The Honorable Chief Minister of Gujarat and the chairman of the State Board for Wildlife has kindly consented to convene a special/urgent meeting as requested by some members to discuss the aforesaid directions and other issues.” The core agenda of the meeting was to chalkout a strategy to take the legal battle ahead against Madhya Pradesh in the Supreme Court.

According to sources who were given the circular to attend the meeting, there were three points on the agenda. But the meeting was specifically called to discuss strong representation in review petition which will be filed within a week. Sources said the first point on the agenda pertained to the SC judgment for translocation of the lions.

“But just when the letters were about to be dispatched, it was realised that the three MLAs who were part of the board would not be considered as legal members as their term got over with the election. As per rules, we have to inform members about the meeting in advance.

To ensure that the meeting is not considered ‘illegal’ the three members were selected immediately and they were sent the letters on Monday itself,” said a SBWL member who is going to attend the meeting. Those who were nominated as members in MLAs category include Moti Vasava from Dediyapada of Narmada district, Shankar Chaudhari from Radhanpur and Kanu Desai from Pardi in Valsad district.

Maharashtra officials get animal rescue training in Gir.

AHMEDABAD: A team of 13 officials from Maharashtra, including chief wildlife warden of Maharashtra SWHNaqvi, visited Gir sanctuary for the three-day study on rescue of wild animals, including leopard.

This was the second such training imparted to forest department officials from Maharashtra in Gir. The first batch had officers who were below the rank of CCF and the present batch had officers of level of additional principal chief conservator of forest and chief conservator of forest. They visited Gir for three days from April 21.

Their visit was a follow up of principal secretary, forest and environment, Maharashtra, Praveen Pardeshi, who visited Gir in March along with the officers of forest department. R L Meena, CCF, wildlife circle, Junagadh; Sandeep Kumar, DCF, wildlife division, Sasan-Gir hosted the training session.

"Gir forest management is becoming a model for study in human-wildlife conflict management in India. Skilled manpower, modern infrastructure, quick responses, community cooperation, strong communication system, community awareness among others are the base for the strength of the successful rescue operations and treatments in Gir protected area," Kumar said.

Maharashtra officials were also surprised to see women involved in the rescue of animals. Rasila Wadher, the only woman in the rescue team, demonstrated how to rescue the animal, treat it and even release it after treatment.

Kuno Wildlife Sanctuary: Ready for lions.

The Asiatic lions of the Gir forest are among the most threatened populations of large carnivores in the world. Photo: AFP
The Asiatic lions of the Gir forest are among the most threatened populations of large carnivores in the world. Photo: AFP

Updated: Tue, Apr 23 2013. 02 15 PM IST
Faiyaz A. Khudsar

Asiatic lions can now be relocated from Gir to the Kuno Wildlife Sanctuary. Faiyaz A. Khudsar says the move has been many years in the making
“Dilli ke Patrakar” (journalist from Delhi). In spite of staying with the Saharia community in a hut (locally known as pataur) for over a year, extending support and help in the relocation and rehabilitation of villagers from within the Kuno Wildlife Sanctuary and of my subsequent, more than 10-year association with Kuno and its neighbourhood communities, this is still how I am locally known.
I first went to Kuno in 1999 with a friend at the invitation of the then divisional forest officer (DFO), J.S. Chauhan, now chief conservator of forests (CCF). In those days, only one bus used to ply between Vijaypur tehsil and Arrod village and I walked seven kilometres on a water-logged and muddy trail to Agraa village to meet him. Today many buses ply between Vijaypur and Agraa.
The Kuno Wildlife Sanctuary is situated in Sheopur district of Madhya Pradesh. The total area of 1,269 square kilometres (sq. km) is now managed as the Kuno Wildlife Division, Sheopur. That includes about 924 sq. km of the surrounding forest habitat of Sheopur Forest Division that has been brought under the Kuno Wildlife Division, besides the sanctuary area of 344.686 sq. km. These changes were made to prepare the area for the introduction of a new population of lions.
The sanctuary derives its name from the Kuno, a tributary of the river Chambal; this perennial river flows through the middle, bisecting the sanctuary. The sanctuary was notified as the Kuno Wildlife Sanctuary in 1981, but it is popularly known as the Kuno Palpur Sanctuary owing to a seventh century fort in Palpur, situated on the banks of the river Kuno. This used to be the capital of the Palpur jagir under the Scindia dynasty of Gwalior. Palpur is one of the biggest evacuated villages within the sanctuary and there is an old forest rest house on the banks of the river Kuno which has been a very important destination for me: I stayed there almost six years for my research and continue to visit.
Today, free-ranging Asiatic lions (panthera leo persica) are found only in the Gir National Park and Sanctuary and its surroundings. The lion population fell to its lowest number in 1893 when only about 18-20 of the beasts remained. Since then it has shown a remarkable recovery. But if a small population of large carnivores is restricted to a single site, it faces a variety of extinction threats.
So the Asiatic lions of the Gir forest are among the most threatened populations of large carnivores in the world and require immediate measures to ensure their long-term survival. To realize evolutionary potential, survival and to promote genetic vigour, it is desirable that wildlife populations be widely distributed in their former historic, geographical ranges. Thus, the reintroduction concept has been developed to ensure long-term survival of the species by creating a new, free-ranging population in areas from where the species had totally disappeared.
To minimize the extinction threats to Asiatic lions, a survey was conducted to identify potential sites for reintroduction. Several potential sites were surveyed, based on sociological and ecological parameters, and ultimately the Kuno Wildlife Sanctuary was selected as the most suitable for a second, free-ranging population of Asiatic lions.
When the site was selected, the first task was to relocate and rehabilitate 24 villages from within the sanctuary. A cabinet sub-committee of the Madhya Pradesh government visited Palpur village in January 1996 to talk to the villagers, and then over 1,500 families, mainly from the Saharia tribe, were rehabilitated, reducing the biotic pressures drastically. I started habitat monitoring and prey base estimation within the sanctuary after the villagers’ relocation and rehabilitation.
The forest of Kuno is classified as a northern, tropical, dry, deciduous forest and is dominated by species such as Kardhai, Khair, Dhawa, Salai and extensive savannah woodland. The sanctuary supports very rich faunal diversity that includes fish, amphibians, reptiles, birds and mammals. The important prey species include chital, chinkara, sambar, nilgai, four-horned antelope, wild pig and the common langur. Leopard, dhole and grey wolf are the main carnivores; occasionally, a tiger sighting is also reported from the region. Today, Kuno is ready to receive Gujarat’s lions with a very encouraging trend of a prey base that is almost 50 animals per sq. km.
Someday soon, I will huddle in a bed at Palpur guesthouse, listen to the alarm calls of the chital and know that the lions of Kuno are on the move. This is the goal I worked towards—to realize the dream of rehabilitated villagers from the Kuno Wildlife Sanctuary and many staff and researchers.
Faiyaz A. Khudsar has a PhD in biodiversity assessment and prey base estimation in Kuno Wildlife Sanctuary. Kuno Wildlife Sanctuary in Madhya Pradesh is the place that has been chosen for the translocation of lions from Gir in Gujarat. He’s currently a senior scientist at the Yamuna Biodiversity Park in Delhi.

The Cat Women of Gir Forest.

Devika Chaturvedi  April 19, 2013 | UPDATED 10:31 IST
The Cat Women of Gir Forest: Modi's pride of women guards is working wonders in the Asiatic lion's last abode

Raseela Vadher knows no fear. Unafraid, she ventures into Gir forest peering intently through the bush looking for signs of trouble. It is not the presence of wild animals that bothers her. She is on the lookout for poachers, illegal gangs felling prized teak trees, or local tribesmen lighting fires that could set the jungle ablaze.

Remember the young woman forest guard Narendra Modi so proudly described to FICCI'S women entrepreneurs in Delhi on April 8? "More than lions, visitors are amazed at the sight of this gutsy girl who fearlessly walks amid a pride of wild lions," he said.

Just 26 years old and already a veteran of the Gujarat government's extraordinary initiative to protect the only natural abode of the Asiatic lion, Vadher is that young woman. From a poor family of Junagadh's Bhanduri village, she signed up as a forest guard in August 2008 and has by now been out on over a thousand successful missions, including about 350 to rescue lions in distress. She loves the forest and this is her dream job.

Like trophies from battle, she happily wears 15 deep scars from a nearfatal lion attack in the summer of 2012. And she is not alone: There are 40 other women van raksha sahayaks, equally driven and zealously watching over the Gir's precious bounty.

Chief Minister Narendra Modi's inspirational move in 2007 to employ women to guard the reserve forest may well have taken a cue from Gir's lionesses, who relentlessly hunt down the plentiful chinkara, nilgai or spotted deer to feed their pride, and turn fiercely protective when their cubs are threatened. Junagadh's women are proving equally enthusiastic in protecting the sanctuary.

An Asiatic lion at Gir
An Asiatic lion at Gir
At last count (April 2010 census), there were 411 lions in the 1,412 sq km of reserve forest-52 more than the last count in 2005. There's been more good news since Vadher and her colleagues joined. In November 2010, GPS monitoring of the big cats showed that as many as 50 adult lionesses were pregnant. Though surviving cubs from the births in early 2011 will only be counted in the next census in 2015, wildlife officials are confident "the pride will have grown significantly". The Divisional Forest Officer (DFO) at Gir, Sandeep Kumar, acknowledges the impact of the women guards. "The numbers are rising because they (the women guards) have been successful in creating a new awareness amidst women and children in villages near the forest," he says. Assuming a more gently persuasive approach than their male counterparts who worked this beat before, Vadher and her colleagues, like Jayshree Patat, 26, and Shabnam Rinbaloch, 24, have worked hard to win cooperation not just from local villagers but also from maaldharis, the semi-nomadic tribal herdsmen who live in the sanctuary.

Rinbaloch says her job in the sanctuary has been a hugely empowering experience. Belonging to a tribal Muslim community that did not let its womenfolk seek work outside home, she became the trailblazer for other young women in her village of Jamanvada. In 2009, three others joined what they proudly refer to as the 'women's brigade', each bringing home Rs.5,200 a month to add significantly to their meagre family incomes. Thrilled with the overwhelming success of the initial recruitments, Aradhana Sahu, deputy conservator of forests at Junagadh, and DFO Kumar, are preparing to hire 100 more, to buffer preservation and protection initiatives in the Gir forest. Kumar says 1,200 young women have applied to join training as forest guards this year.

Before Modi had his brainwave of recruiting women guards in 2007, the Gujarat Forests Department was a decidedly male bastion with just two women, both Indian Forest Service (IFS) officers, amid its ranks. "It was fascinating to watch them at work," says wildlife enthusiast Madhavendra Singh, who came to see the lions up close in Gir. But quite like what Modi spoke about at the FICCI gathering, the 27-year-old from Bhopal says he was completely stumped by the way the wild beasts appeared almost tame in the presence of the women guards. Tourists and animal lovers are returning to Gir in a flood; many, like Pune's Revati Krishnan, 40, inspired by Amitabh Bachchan's captivating 2011 campaign for Gujarat Tourism Department. In just two years, tourist footfall has doubled to 302,428 in 2012-13.

But for Gir's 'women's brigade', it is more than about getting visitors a decent photo opportunity with the big cats. "Preserving the forest is serious business," says 25-year-old Manisha Vaghela, who singlehandedly tracked down and apprehended a gang of nine motorcycle-borne poachers trying to hunt chinkara antelopes in 2011.

Armed with double-barrelled shotguns and walkie-talkie radios, the vigilant women fan out into the bush, unmindful of the danger-venomous snakes, crocodiles lurking around watering holes, a hungry leopard or angry lionesses protecting their cubs. On an average day, each of the women's patrols negotiates over 25 km of forest, even during peak summer months when temperatures inside the sanctuary top 45ºC.

Vilas Antana, 24, graduated in Sanskrit from a college in Amreli and knew nothing of wild animals till the day she signed up to work as a guard in Gir. As with many of her colleagues, the job has diluted her marital prospects, with few young men of her community willing to marry a woman who wears khaki and wields a shotgun. She is not overly bothered, though, happy instead with the independence her job provides. "I can now tell you scientific names of all the birds and animals in this forest," she says.

Gir lions: Gujarat to study SC verdict before taking next step.

Press Trust of India | Updated Apr 16, 2013 at 10:06am IST
Ahmedabad: The Gujarat government on Monday said it will study the full judgement of the Supreme Court on relocation of Asiatic lions to Madhya Pradesh before deciding its future course of action on the issue. Currently, Gujarat's Gir sanctuary is the only home to Asiatic lions, an endangered species.
The apex court on Monday directed the Centre to shift the Asiatic lions from the Gir forest to Kuno Sanctuary in Madhya Pradesh, saying the big cats need a second home to prevent their extinction due to some epidemic or large forest fire. "Before taking any step, we will study the Supreme Court judgement to find out on what basis they have arrived at this decision," Gujarat Finance Minister and Government spokesperson Nitin Patel said.
"The Gujarat Government has maintained that relocation of lions outside Gujarat is not desirable as it is an emotional issue for the people of the state, whose efforts over the years have led to manifold increase in the number of the big cats in the Gir wildlife sanctuary," he said. A century ago, Gir housed around 100 Asiatic lions, whose number, as per the last census conducted in 2010, stood at over 400.
Gujarat to study SC verdict on lions before taking next stepThe apex court on Monday directed the Centre to shift the Asiatic lions from the Gir forest to Kuno sanctuary in Madhya Pradesh.
The State has been opposing the relocation of lions, considered as "pride of Gujarat", to Kuno sanctuary. Meanwhile, Rajya Sabha MP Parimal Nathwani said the verdict has disappointed wildlife enthusiasts and experts in the State.
Gir offers unique environment for the growth of Asiatic lions and the new home may not be suitable for them, said Nathwani, who is also Group President (Corporate Affairs) at Reliance Industries.

Who will be the lion king? Shivraj Singh Chouhan challenges NaMo for pride of place.

Apr 20, 2013, 12.00AM IST
The collective word for lions is 'pride'. And never has this word been more appropriate than in the current context of the controversy created about some lions from the Gir forest of Gujarat being relocated, thanks to a Supreme Court order, to Madhya Pradesh - a move which appears to have stung the 'asmita' of the state presided over by Narendra Modi.
The proposal to shift a few of the Gir lions is based on the observations made by an environmentalist's report, which has subsequently been upheld by the apex court, that as it is deemed unwise to keep all one's eggs in one basket, it is also imprudent to keep all one's lions in a single forest. As of now, the endangered Asiatic lion has only one habitat left, namely Gir. Were an epidemic to strike the area, the leonine species could become extinct, at least in the wild.

Gir forest officials and other local residents have opposed the move, arguing that the badlands of Madhya Pradesh, rife with poachers and dacoits and short on natural prey for carnivores, are far from ideal for the resettlement of Gujarat's lions. However, there could be reasons other than environmental behind such resistance, which could be influenced by the compulsions of political symbolism. In popular lore, the lion is considered to be the king of the jungle. With his openly aired prime ministerial ambitions, Gujarat CM Narendra Modi is being projected by his supporters as being the future king of India's political jungle.
Coincidentally enough, Madhya Pradesh chief minister Shivraj Singh Chouhan is seen by some in the BJP to be a serious rival to NaMo as the party's candidate for the PM's gaddi after the next general elections. Does the relocation of the Gir animals contain a political subtext, that ShiCho might outdo NaMo in grabbing the lion's share of intra-party support for the candidacy of the prime ministerial post?
Such suspicions might not be entirely baseless. Even as the king of the beasts in the wilderness must contend with adversaries in the form of wily shikaris, the political lion of Gujarat has found himself a formidable foe in Bihar CM Nitish Kumar, who has threatened to have his JD(U) quit the BJP-led NDA combine if NaMo is nominated as its PM-to-be.
Has the Gujarati lion met his match in the up-and-coming contender from Madhya Pradesh, not to mention the Bihari big game hunter who seems to have set his sights squarely on NaMo? It's said that pride goes before a fall, a saying which may hold true for lions of all varieties, zoological as well as political. But do watch this space as more thrilling episodes in this saga of contending lion kings unfold.

Watchtowers to deter poachers at Nalsarovar.

Himanshu Kaushik, TNN Apr 24, 2013, 05.12AM IST
AHMEDABAD: Now on, the watchful eyes of foresters will be trained on the winged visitors from overseas that flock to Nalsarovar every year. The state forest department has decided to set up watchtowers near the bank in three villages where poaching has been rampant. Officials have also decided to allocate special funds to set up an information network for Nalsarovar, a Ramsar site.
Officials in Nalsarovar said that at meetings between senior officials on Monday, it was decided to have watchtowers in three sensitive villages - Darji, Ranagadh and Vekariya. These have sizeable populations and notorious for the poaching of birds. In all, about 9000 people in these villages hunt and eat wild animals.
The meeting was held after two people from Ranagadh village were caught with 60 coots that had been trapped and whose legs and wings were broken to keep them from escaping.
These watchtowers will be equipped with closed circuit cameras and in order to prevent damage, one hidden camera will also be installed to catch poachers who will manage to find the visible ones.
Sources in the department said that against the sanctioned strength of 12, there are just six staffers - including the deputy conservator of forests and assistant conservator of forests. With just half the sanctioned strength, officials have to keep watch over 1.5 lakh winged visitors and thousands of tourists to the wetland spread over 120.82 sq km.
"There are only three guards to secure the highly sensitive site, which has communities known for poaching living nearby," said a source. It was decided at the meeting that staff strength would be increased to ensure illegal activities are monitored efficiently.
Secret fund underway
Ahmedabad: It takes a horrific incident of poaching for the state government to loosen its purse strings. Unlike the police department, the forest department did not have a provision for any such secret funding. The poaching of 60 coots was enough for the state to allocate funds for Nalsarovar. An information network will be created with that money.
It was after the poaching of eight lions in April 2007 that the state government allocated funds for Gir Sanctuary. Officials said that a special secret fund will also be set up for Thol. "There is a need for such special funds in all the sanctuaries, but the government rarely makes any provisions. These will help officials keep a check on not just poaching, but also on the movement of suspicious persons.
Not surrendered yet
Forest officials have arrested Kadar Khamasa and Valli Mohammed, residents of Jambu village in Limbdi taluka of Surendranagar, but have been unable to trace those who supplied coots to the duo. The two had applied for bail but their pleas were rejected. This in turn, proved a deterrent from surrender for the others. The remaining four accused had promised to surrender but once bail was denied, they have now gone missing along with their families.

'Human assisted dispersal' of lions a good move: Bombay Natural Hisotry Society.

Linah Baliga, TNN Apr 21, 2013, 05.56PM IST
MUMBAI: The Supreme Court of India's recent judgment permitting translocation of some of the endangered Asiatic lions from Gujarat's Gir National Park to Kuno Palpur Wildlife Sanctuary in Madhya Pradesh is considered to be a good and essential step for the long-term survival of the species.
The Bombay Natural Hisotry Society(BNHS) India, terms this type of re-introduction as 'human assisted dispersal', which means re-introduction of a species to some part of its former range through human intervention. BNHS is of the opinion that many other threatened species can be conserved using this approach wherever it is necessary and appropriate to do so.

BNHS observes that whenever natural dispersal of wild species is not possible any longer due to lack of habitat corridors because of human activities and settlements, it is essential to have 'human assisted dispersal'. Dr Rahmani elaborates on the point saying that although there has been good growth in the numbers of Asiatic lions in Gujarat following conservation measures, there are no forest corridors available at present for the animals to disperse to other areas of their former range in other states. In such cases 'human assisted d'ispersal is required. The same can be used for other threatened species on case to case basis.


Sunday , April 21 , 2013

The Gir lion is emblematic not just of the state of Gujarat. It is also the pride of India. The lions in the Gir National Park and Wildlife Sanctuary are the last of the species called panthera leo persica and there are only around 400 of them left. For decades now these lions have been nurtured and looked after in the Gir National Park. The lion is not indigenous to India. According to one theory, lions from Persia and Africa first started being imported into India some 2000 years ago; the import increased with the demand for lions among Indian royals. It should be mentioned in contradistinction to the above theory that there are differences between the African lion and those found in Gir. The latter has a sparser mane, for example. The breeding of the Gir lion in India began in the 19th century. This was done under the direct patronage of the nawab of Junagadh under whose territory fell large tracts of the Gir forest. Apart from their dwindling numbers — in fact, they were retrieved from near extinction by the aforementioned nawab and the Gujarat state government — and their uniqueness there are certain other features that make the lions in the Gir National Park very distinctive. Observers have noted that the lions in the park are remarkably tame. This is probably related to the fact that for decades the lions have been intensely managed by human beings. They often behave like giant dogs and ignore the available cover. All these factors suggest that these lions need to be preserved.
It is, however, increasingly becoming a difficult proposition to preserve the lions in the Gir National Park. For one thing, the sanctuary is no longer large enough for the lion population and the lions are tending to move away from the river towards the sea. Further, it does not make sense to keep the entire species in one enclosed space. An epidemic attacking the lions or a natural disaster affecting the area could wipe out the population or a very large part of it. Environmentalists have been voicing their concern about this for a long time. Finally, a solution has been found. Experts in the Wildlife Institute of India have recommended that some of the lions should be relocated in the Kuno Wildlife Sanctuary, which, in terms of vegetation and cover, is best suited to be a habitat for lions. The Supreme Court of India has upheld the recommendation and directed that the lions be relocated. This is a major step in the preservation of the panthera leo persica.
The fear that because of this relocation Gujarat will lose some of its tourist traffic is unfounded. The Indian rhinoceros is found in forests like Jaldapara and Gorumara but lovers of wildlife still rush to Kaziranga to see the rhino. Similarly, Gir will remain associated with lions. Moving a pride of them will not diminish Gir’s or Gujarat’s pride. Politics is about human beings — keep lions and animals out of or above it.

Pygmy Elephants, Asiatic Lions and Other Links from the Brink.

pygmy elephant baby
April 20, 2013
Pygmy elephants, Asiatic lions and Siamese crocodiles are among the endangered species in the news this week.
Pygmy elephant update: Remember the 14 pygmy elephants (Elephas maximus borneensis) that were poisoned in Borneo back in February? There’s both good and bad news about the case. The good news is that Baby Joe, the youngster that survived the poisoning (and pulled at our heartstrings after he was photographed trying to wake his dead mother) is doing well. The bad news is that the elephants’ exact cause of death still has not been identified, nor has their killers. The investigation, however, continues.
You can take the Gir lions out of Gir but you can’t take the Gir out of Gir lions: India’s Supreme Court this week approved a plan to move a small number of rare Asiatic Lions (Panthera leo persica) from the Gir forest in Gujarat to a new reserve in Madhya Pradesh. This is great news for the lions, which are so beloved and identified with Gir that they are typically referred to as “Gir lions.” There are now too many lions to fit comfortably in Gir (see my article about them here), and keeping them all in one place increases the likelihood of a single disaster affecting the entire population. The new reserve will help to alleviate both situations. In addition, this could help to give the lions room to increase their population level beyond their current 400 or so big cats. The people of Gujarat do love their lions and have long fought the proposed move—and even protested the Supreme Court’s decision this week—but as long as they are protected in their new reserve this is a change that should be welcomed.

siamese crocodiles released
Photo by Alex McWilliam/Wildlife Conservation Society

What a croc (in a good way): Here’s some good news for the critically endangered Siamese crocodile (Crocodylus siamensis). Nineteen baby crocs, each 19 months old, were released into wetlands in Lao PDR this week. They’ll live in a “soft release” pen for a few months to get acclimated to their new habitat, after which they’ll be allowed to go forth and do crocodilian things. Only an estimated 250 Siamese crocodiles remain in the wild today.
Rhino horns and toe nails: WWF and TRAFFIC, the wildlife trade monitoring network, have launched a new ad campaign to convince people in Vietnam not to buy rhino horns. Since rhino horn is made out of keratin, the same stuff that’s found in human fingernails, the clever ads replace a rhino’s horns with an image of feet. Ick. Vietnam has become one of if not the top market for illegal rhino horn, where it is used as a hangover cure, detoxifier and sexual stimulant (it doesn’t really do any of those things).
One of the ads appears below. The copy translates “Rhino horn is made of the same stuff as human nails. Still want some?” Awesome.
rhino horn toe nails

Adding insult to injury: A boat smuggling more than 10,000 pounds of frozen pangolin meat struck and damaged a protected coral reef in the Philippines on April 15. The Chinese smugglers-slash-fishermen on board the vessel are being held and could face still jail time and fines. Good. All pangolin species (also referred to as “spiny anteaters”) are increasingly threatened by poaching and smuggling for use in traditional Asian medicine and for their meat.
Well, that’s it for this time around. For more endangered species news stories throughout the week, read the regular Extinction Countdown articles here at Scientific American, “like” Extinction Countdown on Facebook, or follow me on Twitter.
About the Author: Twice a week, John Platt shines a light on endangered species from all over the globe, exploring not just why they are dying out but also what's being done to rescue them from oblivion. Follow on Twitter @johnrplatt.
The views expressed are those of the author and are not necessarily those of Scientific American.

The lion's homecoming.

Rajat Ghai  April 19, 2013 Last Updated at 21:36 IST

Wildlife enthusiasts will soon be able to see the lion in Madhya Pradesh also. Rajat Ghai traces the big cat's journey in India down the ages.

Once upon a kingdom
From Vedic and Biblical times to the 19th century AD, the Asiatic Lion (Panthera leo persica) roamed over a swathe of Asia, from Turkey in the west to Bihar in the east, and from the Caucusus in the north to the Narmada river in the south. In the Indian subcontinent, the lion lorded over Punjab (Pakistan and India), Sindh, Baluchistan, Haryana, Delhi, Rajasthan, Uttar Pradesh, Gujarat, Madhya Pradesh, Chhattisgarh, Bihar and Jharkhand. Lions in the subcontinent largely inhabited dry, tree-covered Savannah and low-scrub jungle in comparison to the Bengal Tiger which inhabited mature-tree forests.

Ancient India
The lion's impact on South Asian history, culture and art is significant. Mahesh Rangarajan notes in his paper 'From princely symbol to conservation icon: A political history of the lion in India': "It (the lion) was perhaps only rivalled in its power over the human imagination in India by the tiger." Adds Divyabhanusinh Chavda, author of The Story of Asia's Lions: "In the Vedic period, you had Narasimha ("Man-Lion"), the fourth of the Dashavatara of Lord Vishnu. The Buddha was known as 'Shakya Simha', the Lion of the Shakyas. His first sermon at Sarnath has been likened to (and is known as) by Buddhists as 'Simhanada' (Lion Roar). The lion is also the symbol of Mahavira. And we, of course, know of Emperor Ashoka's association with the lion."

According to legend, 2,500 years ago, Vijaya, a disinherited Indian prince migrated to the island of Lanka with 700 followers. Vijaya's grandfather was a lion. His descendants, the Sinhala ('Lion people'), are today the majority ethnic group on the island, the flag of which is emblazoned with a lion too.

Medieval India
From 500 AD, Rajput princes across India started adopting the title Simha instead of the classical Varman. Today, we know this surname as 'Singh', most commonly associated with Rajputs and Sikhs.

Lions find constant reference in the Delhi Sultanate and Mughal periods too. Sher Shah Suri, the Afghan warlord from Bihar who dethroned Humayun, reportedly acquired the name 'Sher' after killing a lion with his bare hands. The Mughals, Persianised Turko-Mongols from Central Asia, used the regal Persian Sher-o-Khurshid ('Lion and Sun') as their personal coat-of-arms.

The Leonine Holocaust
Lions in India were never maneaters like their African cousins, but were notorious cattle thieves. They were also hunted for sport. "Colonel James Skinner (1778 -1841), the famous founder of Skinner's Horse and builder of Delhi's St James' Church, is recorded as shooting lions on horseback. Another officer, Andrew Fraser killed 84 lions," says Chavda. "In 1810, a General Mundy shot a lion near Hansi (modern Haryana). In the first half of the 1800s, British soldiers stationed at a cantonment in Deesa (North Gujarat) are recorded as spearing lions. And the all-time record in lion hunting in India goes to George Acland Smith, an officer who shot 300 lions near Delhi in 1857, on the eve of the Sepoy Mutiny," he adds.

Soldiers of Maharaja Ranjit Singh, the Sikh sovereign of the Punjab, are recorded killing lions with bayonets in Lahore in the 1830s. In Patiala, hunting lions was an annual affair in which "400 horsemen beat vast plains teeming with antelope and Nilgai" (Rangarajan).

The End
All this killing eventually did its job. Region after region recorded local extinctions: Bahawalpur (1800), Palamau (1814), Haryana, Baroda and Ahmedabad (1830), Sindh (1842) and Gwalior (1872).

By 1890, when Prince Victor Albert visited India, the only place where he could find lions was the Gir forest in Kathiawar, where politics between three feuding princely states (Junagadh, Bhavnagar and Baroda) and the British Government in Bombay ensured safety for lions. Later, the Nawabs of Junagadh took on the task of preserving lions in Gir, where they survive till today.

Uproar in Sasan against lion relocation, bandh today.

Thursday, Apr 18, 2013, 14:13 IST | Place: Ahmedabad, Gandhinagar | Agency: DNA
Amid demands for review petition, an activist has threatened self-immolation.
The Supreme Court, in its judgment on Gir lions, may have quashed Gujarat’s prejudices by sharing its Pride, but the state seems to be in no mood to let its hurt pride suffer in silence. An agitation is brewing against the verdict on translocation of lions from Gujarat to MP. Local tempers in the sanctuary are rising, so much so that some have also threatened to self-immolate if the committee to be set up on SC directions does not stop the shift.

“This is a very unfortunate decision, which is based on a biased report prepared by scientists.

I’m garnering public support against the decision and fifty of us ready to end our lives through self-immolation,” claimed Dinesh Goswami, president of Kodinar-based Prakruti Nature Club.

But some others believe that filing a review petition is the best option. “We have sent a memorandum demanding review of the case. We want state government to file a review petition in the issue. It is an SC verdict and our hands are tied,” said Bhikhubhai Jethva, president of Khambha-Gir Nature Club.

“The argument under which translocation is approved by the Supreme Court is based on flawed research,” said president of Sasan Gir Hotels Association, Mehul Mehta. He said that there is ample scope of relocation of lions in Gujarat itself.

Meanwhile, traders, hoteliers and forest beat guards are also joining the protest. The area is going to observe a bandh on Thursday under leadership of sarpanch of Sasan and Bhalchel. A protest rally has also been planned on Saturday which will be joined by residents in sanctuary area and villagers.

Govt to file review petition in lion translocation case
Gujarat Government on Wednesday decided to file a review petition in Supreme Court, challenging the apex court’s directive to translocate of Asiatic Lions from Gir Forests to Kuno-Palpur in Madhya Pradesh. SC had on Monday ordered transfers of the lions to MP, creating a political and social stir in Gujarat.

“We have received copy of the judgement and have studied it primarily. The government has decided to fight the legal battle till the last option to ensure that the lions remain in Gujarat,” said Nitin Patel, state finance minister and spokesperson, indicating that the government will be filing a protest petition in the matter. “We have argued in the case, in detail, and filed affidavits about the facts on Asiatic Lions, including its genesis to the recent development.”

A senior government official, who has worked in the case, has indicated that it wouldn’t be difficult for the government to stop transfer of lions to MP.

“In one of the affidavits filed, we have written about the lions’ behaviour with locals in Gir,” said the official. “People there love the animal and ensure its safety. The lion is habituated to human presence.”

“This kind of peaceful, non-conflicting coexistence may not be possible in Madhya Pradesh,” he added. “Also the area provided for the lion (in Madhya Pradesh) isn’t enough. There is possibility of lion-tiger, lion-human conflicts.”

Pride hurt: Murmurs of protest fill Gir air against lion translocation.

Wednesday, Apr 17, 2013, 13:42 IST | Place: Ahmedabad | Agency: DNA
Forest guards and residents carrying placards and shouting 'Chellam Go Back' protest scientist's entry into Gir.
Even as the Gujarat government mulls its next plan of action regarding translocation of lions to Madhya Pradesh, a few groups in Gir are gearing up to protest Monday’s verdict of the Supreme Court.

On Tuesday, a small group of residents protested the presence of wildlife expert Ravi Chellam in the area. “There had been murmurs of protest, but the biggest outburst came later in the day when Sasan residents realised that one of the scientists – Ravi Chellam – who was a member of the committee that advised translocation of lions, was in the village,” said a resident.

Some residents and a few forest guards began a protest as the scientist tried to enter the sanctuary, he said, adding, “They even carried placards that read ‘Chellam Go Back’ and prevented his entry to the sanctuary. He had to return to the the forest department guest house.”

When contacted, deputy conservator of forests (DCF) of Sasan, Sandeep Kumar, admitted that a protest was carried out. “It was taken out by the local guides and not the forest guards,” said Kumar. He refused to comment further. Meanwhile, activist Bharat Sojitra claimed that the people want the forest department to ensure that herbivores, too, were reduced, if lions were taken. 

“It is because of the large lion population that the herbivores are in control. Once they go, it will be difficult to control them,” said Sojitra. Pointing out that the lions often come to farms, too, he said that it was a blessing in disguise as it helped keep the wild herbivores away and farmers did not have to protect their fields. There is already a rise in herbivore population and moving a large number of lions would only worsen the situation, he added.

Another activist, Dinesh Goswami of Prakruti Nature Club, said that 50 NGOs had submitted a joint memorandum to the collector to help prevent translocation of lions. “We are also spreading awareness about the issue. There will be regular programmes on an everyday basis and we plan to involve school and college kids as well,” said Goswami.
Wednesday, Apr 17, 2013, 13:42 IST | Place: Ahmedabad | Agency: DNA
Even as the Gujarat government mulls its next plan of action regarding translocation of lions to Madhya Pradesh, a few groups in Gir are gearing up to protest Monday’s verdict of the Supreme Court.

On Tuesday, a small group of residents protested the presence of wildlife expert Ravi Chellam in the area. “There had been murmurs of protest, but the biggest outburst came later in the day when Sasan residents realised that one of the scientists – Ravi Chellam – who was a member of the committee that advised translocation of lions, was in the village,” said a resident.

Some residents and a few forest guards began a protest as the scientist tried to enter the sanctuary, he said, adding, “They even carried placards that read ‘Chellam Go Back’ and prevented his entry to the sanctuary. He had to return to the the forest department guest house.”

When contacted, deputy conservator of forests (DCF) of Sasan, Sandeep Kumar, admitted that a protest was carried out. “It was taken out by the local guides and not the forest guards,” said Kumar. He refused to comment further. Meanwhile, activist Bharat Sojitra claimed that the people want the forest department to ensure that herbivores, too, were reduced, if lions were taken. 

“It is because of the large lion population that the herbivores are in control. Once they go, it will be difficult to control them,” said Sojitra. Pointing out that the lions often come to farms, too, he said that it was a blessing in disguise as it helped keep the wild herbivores away and farmers did not have to protect their fields. There is already a rise in herbivore population and moving a large number of lions would only worsen the situation, he added.

Another activist, Dinesh Goswami of Prakruti Nature Club, said that 50 NGOs had submitted a joint memorandum to the collector to help prevent translocation of lions. “We are also spreading awareness about the issue. There will be regular programmes on an everyday basis and we plan to involve school and college kids as well,” said Goswami.

Forest Rights Act extended to non-tribal areas.

Express news service : Ahmedabad, Fri Apr 05 2013, 04:18 hrs
The state's Tribal Development Department (TDD) has extended the Forest Rights Act to non-tribal regions of Gujarat, paving the way for forest dwellers such as maldharis (pastoralists) to legally lay claim over areas they inhabit. Some officials predict that it would force other departments to act especially in terms of halting talked-about maldhari evictions in Gir, Saurashtra and revenue settlements in the Banni Grasslands, Kutch.
The Forest Rights Act has been in force in the state for more than five years now, although it has been restricted to 12 districts covering 43 talukas in the eastern tribal belt.
The government resolution issued by the TDD as Assembly elections approached last year but which was apparently kept under wraps due to the model code of conduct, says district collectors would be chairpersons of district-level committees formed under the Act, with three members of each district panchayats to be nominated members.
However, posts of member-secretaries would be held by officials of the TDD - the department's planning officer at Talala would be member-secretary of committees in Junagadh, Porbandar, Amreli and Jamnagar.
The department's vigilance officer of Rajkot will hold the post for Rajkot, Surendranagar and Bhavnagar, while his counterparts in Ahmedabad would be member-secretary for Ahmedabad, Anand, Kheda and Gandhinagar and the one in Palanpur for Patan and Mehsana. The Kutch social welfare officer would be member-secretary for Kutch region.
As for expected changes in the Gir forest, senior forest officials said the "worrisome" increase in domestic animal populations would now have to be tackled through means other than eviction, which has anyway come in for criticism lately due to evidence that most of those shifted out in the early 1980s eventually shifted back inside and either sold land allotted to them or converted it for other purposes other than traditional means of livelihood.

Forest staff to get arms training to prevent poaching.

Express news service : Ahmedabad, Sat Apr 20 2013, 05:12 hrs
After the recent bird-poaching case at the Nal Sarovar Bird Sanctuary, the forest department has scheduled arms-training programme for the forest personnel here in the first week of May. The personnel will receive training from the Surendranagar police.
Nal Sarovar is Gujarat's only Ramsar site. Ramsar sites are wetlands of international importance, designated under the Ramsar Convention, an international agreement signed in Ramsar (Iran) in 1971, which provides for the conservation and good use of wetlands.
"The watchmen carry knives, foresters and beat-guards carry 12-bore guns and range officers carry revolvers. But we want to train them to use these weapons," said S Sasikumar, Deputy Conservator of Forests at Nal Sarovar.
Meanwhile, the 120-sq-km sanctuary has been allotted six new foresters from neighbouring divisions to supplement the two foresters and two beat-guards already posted there. The division has also been permitted to hire another vehicle. As of now, just one official vehicle was allotted for officers there.
A proposal to install heat-sensitive cameras in three villages to monitor any future poaching cases has also been submitted to the headquarters in Gandhinagar.
State plans AC tents for tourists
The state tourism department is planning to erect 500 air-conditioned (AC) tents in top 10 tourist destinations in view of the rising number of visitors. The state attracted 22.3 million tourists during 2011-12 and is expecting a 15 per cent increase in tourist arrivals this year.
Though a list of these spots has not been finalised, destinations like Gir, Saputara, Madhupura, Mandvi and pilgrim towns like Ambaji would likely be in the list, sources said. "We are still in the process of finalising the spots for installation of the tents," said Nirav Munshi, manager of Hotel, Tours and Travels at Tourism Corporation of Gujarat Ltd.

Is shfiting of Gir lions to Sheopur dangerous?

Apr 20, 2013, 04.56AM ISTIf the Union ministry of forest and environment is concerned about the African cheetah being gunned down in Sheopur (MP), why is it not showing the same concern for the Asiatic lion? Any forester would tell you it is easier to kill a lion than a cheetah with a gun. The lion lives in perfect harmony with people of Saurashtra who share the habitat with all living beings. Gujarat should argue in a review petition before the Supreme Court that the shifting move is as good as sending the lions before a firing squad.

Villagers hope Amitabh Bachchan to come forward to rescue Gir Lions.

DNA | Apr 19, 2013, 05:01AM IST
Ahmedabad: Villagers in Sasan are hoping that brand ambassador of state’s tourism, Amitabh Bachchan, will come to the rescue of Gir Lions. They want the superstar to take up the cause of translocation of lions.
“We will urge Amitabh Bachchan to take part in our movement to prevent translocation of lions to Madhya Pradesh,” said Laxman Dhokiya, Sarpanch of Sasan.
 “We will urge him to use social networking websites like Twitter and Facebook to take up the cause. He will be able to reach many more people to sensitise them about why lions shouldn’t be moved.”
Five villages of Sasan, meanwhile, observed a bandh on Thursday to protest the translocation of lions. Groups of concerned citizens gathered at several places to submit a written memorandum to the mamlatdar.
The bandh was supported by Gypsy Association, Forest Guides Association and Gir Hotels Association. Head of the hotels association, Mukesh Mehta, said that they are with the villagers in the protest. “But we couldn’t close the hotels, as bookings were done well in advance. Concerns about translocation of the lions are genuine,” said Mehta, adding that they are trying to sensitise tourists about the issue. “We have had small stickers and posters on vehicles, so that tourists can also know of the issue.”
Though business in the five villages – Bhatched, Haripur, Sasan, Chitrod and Bhojde – was affected, tourists continued to visit the sanctuary.
Reports, meanwhile, suggest that locals are keen to be party to the review petition that the government will file challenging the apex court’s order. “Translocation is Supreme Court’s decision and as such, we can’t go against it. But the government can definitely challenge the order and present its case in a much better way,” said Bhikhubhai Jethva, Khambha Nature Club.

Tigers moving from Rajasthan to Madhya Pradesh, officials concerned.

P Naveen, TNN Apr 19, 2013, 03.01AM ISGWALIOR: Movement of tigers from Rajasthan's Ranthambore to the Chambal range in Madhya Pradesh has become a cause for concerned of the two states.
Three tigers who went missing from Ranthambore have been located in Madhya Pradesh including one in Seoda range in Datia district - a mix of reserve and protected forest area and two in Kuno Wildlife Sanctuary, selected for relocating lions from Gir in Gujarat.
After writing to the Madhya Pradesh forest officials to check whereabouts of the missing tigers, a team of officers from Ranthambore visited Datia district last week. They installed camera traps to check its movements. The tiger was finally caught in the camera traps of Madhya Pradesh forest department.
"Our team followed the tiger all the way to MP. We are happy that it's safe there and the officers were very cooperative. We cannot stop dispersal, only concern is its safety," said Y K Sahu, district forest officer Ranthambore.
The tiger in Datia is said to be a 3-year-old cub of Ranthambore's T-26 tigress. Information collected from the forest officials and the GPS tracking by WWF India- Western India Tiger landscape team indicates this cub travelled more than 220 km to reach the Seoda range.
On March 14, 2013 the tiger was found in Seoda range of Datia territorial forest division, which is a forest patch of 55km in length and width of 11-12 km. The range has both reserve and protected forests with the Sindh river flowing in the middle and the Vindhya hill ranges on the western side. There are many villages on both sides
From March 14 till date several samples and pugmarks have been recorded while following the tiger. There was no tiger sightings reported from Datia till one was shot dead by poachers in 1998. Prior to that one was hunted 'legally' by member of a Royal family in 1960, said sources.
"We are very concerned about the tiger's safety. Additional patrolling is being done to keep tab on its movement," said chief conservator of forest (Gwalior circle) S P Rayal.
Kuno is now a house to two Ranthambore tigers, T-38 that was spotted there since last year and other one that moved in recently. Officers had been visiting the place to track the tigers. "We have not received any pictures of the tiger so far. Search is on," Sahu said.

'King of cool' roars into Dublin Zoo with romance on his mind.

Staff at Dublin Zoo hope their new Asiatic lion, Kumar, will strike up a romance
DUBLIN Zoo is hopeful that its latest addition – a rare Asiatic lion – will father the first pride of lions at the zoo in decades.
Seven-year-old Kumar, who was born in captivity at Edinburgh Zoo, has already fathered several cubs at his most recent home in Rotterdam Zoo.
Zookeepers at Dublin Zoo are confident that he will get down to business with the zoo's lioness sisters Zuri and Sita and father the next generation of lion cubs as part of its international breeding programme for endangered species.
Judging by his keen paternal instincts that were revealed after he sired three litters of cubs at Rotterdam, he would make an excellent addition to the breeding programme, said zookeeper Ciaran McMahon.
"He's very socially well adjusted to the two females and we hope he'll produce two litters," he told the Irish Independent.
The last lion cubs born at the Dublin Zoo was back in 1989 and if Kumar mates successfully, he will produce the zoo's first-ever pride of Asiatic lions.
He has already asserted his dominance over the females with his menacing roar, which is a good sign that he will start mating, Mr McMahon said.
Aside from his potential breeding prowess, he's a lovely cat to watch, he added.
"He is a very proud and majestic animal but he's also very laid back," he said.
"He's the king of cool," he said of Kumar's regal pose as he lies on his haunches taking in the sights of the Phoenix Park.
Kumar arrived at the zoo in late March and after a period of quarantine, has been introduced to the Asiatic lions den – which is mirrored after the Gir National Forest and Wildlife Sanctuary in Gujaret, India.
It was established in the early 1900s as a protected area for Asiatic lions whose numbers had been decimated to just 15 in the world due to trophy hunting.
Today there are still only about 350 Asiatic lions left in the wild – with the majority living within the sanctuary.
If Kumar mates successfully, it will take between 105 to 110 days' gestation for a litter to be born, which typically number between three and eight cubs.
A new litter would not only be a welcome addition to the breeding programme, it would also be a boost to staff who are still mourning the loss of Shelia, the last African lion at the zoo who died of old age last August.

Good weather, plenty of prey for lions in Kuno, says Madhya Pradesh.

P Naveen, TNN Apr 19, 2013, 05.30AM IST
SHEOPUR (KUNO-PALPUR): Gujarat, miffed by the court order on its Gir lions, is asking a string of questions regarding the suitability of MP to house the big cats. Among other things, they say both weather and scarcity of prey in the sanctuary at Kuno would pose a problem. Independent MP Parimal Nathwani from Gujarat especially, had expressed reservations about the court's direction claiming that the lions would be forced to live in a new geographical area, where everything from climate, land, water and their food pattern would change.However, officials at the sanctuary -- which TOI visited few days back -- claim that these factors won't be an issue for the lions. Citing a recent survey by Wildlife Institute of India, World Wide Fund for Nature, India and the state forest department, which was submitted to the apex court, officials point out that prey base (sambar, nilgai, wild pig, chinkara, langur, peafowl and feral cattle) in the area has increased 8-fold between 2006-2012. "This is more than the density in Gir or any other park across the country," says Ashok Mishra, former DFO of the sanctuary. Weather, too, won't make a difference, adds another official.
Incidentally, the move to relocate the lions to Kuno is an old one. The process began after a country-wide survey in 1993-94 by the Indian Wildlife Institute and relocation work started in '96- '97. Between 1997 and 2003, 24 villages, comprising 1545 families, were relocated outside the sanctuary to facilitate the process of transfer of the big cats. And now, when the lions seem set to move in, a number of habitat improvement and prey-based works have been undertaken in the sanctuary which is named after the Kuno River, one of the major tributaries of Chambal that flows through the area.
Interestingly, in their new home, the lions would be in proximity with another member of the cat family. The south-western portion of this landscape is patchily connected to Panna Tiger Reserve through the Shivpuri forest area. On the north-western side, this forest region is contiguous with Ranthambore Tiger Reserve across the river Chambal.

SC orders shifting of some Gujarat lions to MP jungle.

Mahesh Trivedi / 16 April 2013

AHMEDABAD - Brushing aside the Gujarat government’s blatant refusal to shift some of its 411 rare Asiatic lions to neighbouring Madhya Pradesh, the Supreme Court on Monday ordered the Centre to translocate the endangered animals within six months.
The number of big cats to be relocated from their only abode in the Gir forests to their new home in the Kuno wildlife sanctuary will be decided by a committee of wildlife experts.
“The species which is on the verge of extinction needs a second home,” the apex court ruled responding to a public interest litigation filed by an environmental group which favoured the shifting to avoid extinction of the extremely endangered species in case of a calamity, a disease or a disaster like fire.
At least 92 lions have died in the past two years in Gir. Intense conservation efforts by the Gujarat government over the past 50 years have brought them back from the brink of extinction.
The two BJP-ruled states have been at loggerheads over the translocation of the lions. Gujarat has been arguing that the Kuno reserve is not suitable for lions as it also houses tigers, and its weather and environment are also not conducive for survival of the majestic animals listed as endangered by the International Union of Conservation of Nature.
While the Narendra Modi government was still studying the verdict, BJP leader Jagdish Bhavsar told Khaleej Times that Madhya Pradesh had failed to protect its own tigers in Panna sanctuary and the lions from the Gir forest might meet the same fate. “There’s no guarantee that lions will adopt Kuno as their new home. Punjab had tried a lion safari project with zoo lions, but all lions died one by one”, he said.
But Madhya Pradesh, which has been making desperate attempts to acquire a few of Gir’s lions to boost tourism in the name of conservation, has claimed that it is equipped with all the necessary infrastructure, expertise and environment for translocating the lions to its own sanctuary.

Bengal Tiger, Asiatic lion and African cheetah: Rajasthan to have all three big cats.

DNA | Apr 16, 2013, 07:59AM IST
Jaipur: The apex court’s decision has rekindled the hope of Rajasthan being honoured by a sacred distinction of being the first state to be a home to all the three Panthera species of the subcontinent. Though, its plan to develop Cheetah Park stands shelved after SC’s refusal to allow African cheetahs from being brought to the country. 
More than fifteen years back, the Rajasthan government had tried to settle Panthera leo Persica, better known as Asiatic Lions in the state’s sanctuaries. In 1996-97, Government of India (GOI) started a program to identify potential areas for translocation of these lions from Gir to some other forest area. 
This was done to ensure that the already endangered population, which was confined only to Gir forest, does not get wiped out due to Tsunami or any disease. Kamal Nath, who was the then minister of environment and forests identified Kuno-Palpur area, on border of Rajasthan and Madhya Pradesh as a potential site for translocation. 
In the following year (1998), the Rajasthan government also constituted a committee of experts which studies the animals at Gir and submitted a report. The report, which was handed over to GOI, narrated that Rajasthan has a better potential than Kuno-Palpur. “The committee asked the GOI to review the choice of site. It mentioned Darrah sanctuary near Kota and Sitamata sanctuary near Chittorgarh as potential sites, since the habitat and prey base was favorable for population expansion” said Rajpal Singh, member- State board for wildlife and a member of the state committee for relocation of lion in Rajasthan.
Interestingly, Asiatic lions were found in the Kota-Bundi region (Darrah sanctuary area) during the British Raj also. Historically, the area was a preffered ground for royal families to shoot lions and tigers. Recently Darrah sanctuary was included in project tiger area but the government can rethink over introducing lions in the area. “It will be a great boon for the wildlife and for the state also if Rajasthan gets its share of lions” said Singh.

They risk life to save forest.

Swati Bhan, Ahmedabad, April 14, 2013
Women forest guards not only fight wild animals but also superstitious beliefs.
The sun was setting peacefully a few months ago. Anita Raval, in her twenties, was returning home after a hard day's work and was on the periphery of the forest.

What happened then benumbed her and literally stopped dead in her tracks. It was just a hand-shaking distance and a leopard majestically walked past her. She regained her guts and immediately alerted her colleagues and managed to drive back the beast into the forest with the help of villagers.

Young Rasilaben Wadher also had a hair-raising experience which will send shivers down the spine of most people. But she is made of a sterner stuff and handled the crisis with elan. “Once I had gone with my team to rescue an injured lion. It almost
attacked us. We kept our cool and slowly returned to our vehicle. The lion slowly retreated into the forest,” she recalled.

Most women, working as forest guards in the last few years, have many such experiences to share. Some bone-chilling stories that would make the weak look for a better place to work. But these women are made of sterner stuff and no wonder, they continue to guard the forests, facing all odds.

It is no surprise that Gujarat Chief Minister Narendra Modi, while addressing an elite crowd of women in New Delhi, cited several examples of fair sex taking risks to excel in society. When he spoke of women achievers, who rose from a modest backgro­und and carved a niche for themselves, he did not forget to make a mention about the tribal women protecting the forests of Sasan Gir that attracts thousands of tourists for a glimpse of the Asiatic Lions.

Their mention could be for obvious reasons—they have been doing a decent job in the last few years. Their recruitment in 2007 made one more male bastion crumble in the state of Gujarat. In 2007, the Department, for the first time, announced that it would recruit women as forest guards, considered to be a challenging and dangerous job by many.

On selection, the women were given practical as well as theoretical training at the Gujarat Forest Rangers College in Rajpipla. “We even trained them in use of arms and on legal matters,” recalled Sandeep Kumar of Gir West Division, Junagadh.  Most of these women belong to Junagadh and have volunteered for the first all-woman patrol in Gir Wildlife Sanctuary—Van Raksha Sahayak.

One such Sahayak, Trupti comes from the small village of Ambalgadh in Talal taluk of Junagadh district. As a Van Raksha Sahayak, her job involves intense field work in the wild. All recruits conduct night patrolling and rescue missions apart from
other regular tasks.

The recruits went through a gruelling test involving an 800-metre run, high jump, long jump and a 15-km observation walk through the jungle. Those selected were imparted training for 10 days. “I was born and brought up in the Gir area. My parents taught me to love and respect wildlife. I am happy to serve here and don’t need another job,” Trupti added.

The recruits come from Maliya, Una, Kodinar and Junagadh taluks. Women beat guards have been appointed at other sanctuaries in Gujarat too, but Gir is a special case since it is the only abode of the Asiatic lion. Rasilaben Wadher said, “I have taken up the duties of a forester in the Sasan Gir sanctuary division and am loving it,” she added.

It is love of the wild that unites all these women. Keyuri Khambda, who joined the forest department in Gujarat recently, opted for the job because of her love for forest and concern for the wild. “I work with the assistant conservator of forests on patrolling and raids,” she said.

The women hold different profiles. Those on the Mobile Squad patrol the entire sanctuary area, including Gir east, west , Sasan and Junagadh district. The Rescue Squad is responsible for the rescue of wild animals in distress. The Wireless Squad coordinates between all the squads. The patrol squad also manages the tourist circuit at Sinh Sadan in Sasan.

Their work includes preventing smugglers from  cutting trees, preventing forest fire and chasing away leopards. Anita Raval, one such beat guard, manning the Shoolpaneshwar Wildlife Sanctuary in Narmada’s Nandod taluk, admits that it is not a fixed time job like 10 am to 5 pm work. “Many a time when fire breaks out in the forest, I have to rush even at midnight,’’ said Raval.

She admitted that she has to venture out alone in the forests, some times just armed with a stick and not only fight animals but battle superstitions of the tribals. Raval is one of the eight beat guards of the total 129 beat guards in Narmada district. “Cutting forest wood is not allowed, though we allow villagers to collect dead wood,’’ says Raval.
She said when they find somebody cutting trees, they do inform the range officer. “We also prevent villagers from lighting a fire in the forest which most of the time they do as a part of a ritual among the tribals,’’ said Raval.

Forest officials in the area added that it was a difficult task indeed to convince tribals not to follow such rituals and foresters were attacked as well for preventing them from practising these rituals. So, many a time under such a situation a team of eight or nine guards go to the scene instead of one beat guard. This is more of a precautionary measure. They also raise nursery in front of their quarters and later transplant them in forest.

The beat guards have also played a vital role in rescuing the animals straying from forests to human habitat. It has been their job to shift these animals to the forest without endangering the life of the animal and humans. The beat guards admit that theirs was indeed a challenging job, but most of them have opted for it as a choice. So, they enjoy the task despite the risks involved.

Emotional protests sweep ground zero.

TNN Apr 17, 2013, 12.34AM IST
AHMEDABAD: The strength of the emotional bond that the people of Gujarat share with the lions was expressed on Tuesday when the residents of Kodinar and Sasan came out on the streets to protest against the translocation of lions, ordered by the Supreme Court. Jeep owners and the Guides' Association were holding a meeting till late on Tuesday night mulling whether to go on strike.

The sarpanch of Sasan has decided to call for a bandh on Wednesday. On Tuesday, when volunteers of the Prakruti Nature club led by Dinesh Goswami and Jignesh Gohil began their protests, even some forest staff -beat guards, jeep drivers and guides - joined the action. Sarpanch Laxman Dhokadia said that as soon as the news spread among the people that the lions would be translocated to Madhya Pradesh, gloom descended on Sasan.
Dhokadia said that even several leaders from Sasan met forest officials and made a representation that the lions should not be translocated. He said that the lions were the state's property and not national assets. "We the people of Saurashtra saved the lions," he said. "How can you take them away from us. We will also give a memorandum to the officials on Wednesday to press for our demand to stop translocation."
Members of several nature clubs took out a rally in Kodinar to protest against the translocation. They stated that the translocation would be dangerous for the lions. The members also said that the people who carried out poaching in Gir in 2007 were from Katni in Madhya Pradesh and hence the lions could not be safe in that state. The clubs' members have also threatened to resort to self-immolation if the lions were shifted to Madhya Pradesh.

In forests, women guards fight fire, leopards alike.

Rajpipla, Narmada, Fri Apr 12 2013, 04:33 hrs
When Chief Minister Narendra Modi talked about women guards of the Gir forest taking tourists to view lions, at a FICCI meet in New Delhi on Monday, he could have as well meant Anita Raval, a forest beat guard working in the arduous terrain of the Shoolpaneshwar Wildlife Sanctuary around the Karjan Dam in Narmada's Nandod taluka, tracking down people cutting trees, preventing forest fire and chasing away leopards, an example that encapsulates the spirit of women in the state.
Raval, 24, is two years into the job as Van Raksha Sahayak. Her work is not time-bound. In the event of fire breaking out in the forest at night, she has to rush to the forest to help douse it so it does not spread. Many a time, she has to venture out into the forest alone, armed with a stick, to persuade tribals not to cut trees or light fire in the forest which they do as part of a ritual. She also plants trees in the forest from saplings that she grows outside of her quarters in Jeetgarh village near the Karjan Dam.
Raval is one of only eight women beat guards out of a total 129 beat guards in Narmada district, according to a forest department data. Her beat is to cover forest falling under Mota Raipur and Jeetgarh villages along the Karjan Dam, which covers a 1,200-hectare area. She hails from Netrang in Bharuch district, situated some 50 kilometres from where she is posted. "I had applied for the job after I saw an advertisement in a newspaper, though I was not sure if I will get selected. I like my job because I love forest, trees, animals," she said.
She has brought along her parents to stay with her. "Sometimes we have to trek 15 kilometres in a day, and have to rush into the forest at night in the event of fire. We take five to seven persons along when we come to know about fire in the forest or people cutting trees," she said.
"Cutting forest wood is not allowed, though we allow villagers to collect dry wood. When we find somebody cutting trees, we inform the range officer. We also prevent villagers from burning forest which they do as a part of their ritual, but which harms the forest no end. We persuade villagers not to do such a thing, but if they do, we try to douse fire so it does not spread," she said. "We have been trained how to douse the forest fire by creating fire line so we could prevent it from spreading. Among other duty, we grow plant in a nursery near our quarter and shift them into the forest when they grow up," she said.
Raval said in the last two years, she helped release at least six leopards in the forest that were rescued from areas with human population. "I once saw a leopard cross the road very close to me as I was passing through the jungle but it passed by peacefully, as I halted on my way for a moment," she chuckled.

Gujarat will have to keep sending its lions to Madhya Pradesh.

AHMEDABAD: The State Board of Wildlife is meeting on Thursday to prepare a strong review petition to be filed in the Supreme Court shortly to oppose the decision earlier this month to shift some Gir lions to Palpur Kuno sanctuary in Madhya Pradesh.

Studying the decision carefully, the authorities here have realized that this is not a one-time transfer of lions. The SC decision states that the gene pool in MP would be kept healthy by sending fresh male lions from Gujarat every three to five years. The wildlife board and the forest department feel that sending males from Gir to Kuno will disturb the social fabric of the prides.

Their main argument, according to top forest officials, will be that the gene pool of Gir lions has improved and not deteriorated as suggested in the project report for translocation. Gujarat forest officials will quote a study — 'Genetic variation in Asiatic lions and Indian tigers' — jointly conducted by the Center for Cellular and Molecular Biology, Hyderabad; Zoological Survey of India, Kolkata; and Center for DNA Fingerprinting and Diagnostics, Hyderabad.

Quoting the study, Gujarat will try to establish before SC that the lions are healthy and have no genetic disorder at present and are not likely to suffer any kind of such disorder in the next several years to come. Forest officials said that they have several scientific evidences to drive home the fact that the health of the lions is steadily improving. Earlier, the litter size was just one or at the most two, it is now normal to see a litter of two or three or even four cubs.

Wednesday, April 24, 2013

The Evolutionary Dynamics of the Lion Panthera leo Revealed by Host and Viral Population Genomics .

PLOS Genetics 
Research Article


The lion Panthera leo is one of the world's most charismatic carnivores and is one of Africa's key predators. Here, we used a large dataset from 357 lions comprehending 1.13 megabases of sequence data and genotypes from 22 microsatellite loci to characterize its recent evolutionary history. Patterns of molecular genetic variation in multiple maternal (mtDNA), paternal (Y-chromosome), and biparental nuclear (nDNA) genetic markers were compared with patterns of sequence and subtype variation of the lion feline immunodeficiency virus (FIVPle), a lentivirus analogous to human immunodeficiency virus (HIV). In spite of the ability of lions to disperse long distances, patterns of lion genetic diversity suggest substantial population subdivision (mtDNA ΦST = 0.92; nDNA FST = 0.18), and reduced gene flow, which, along with large differences in sero-prevalence of six distinct FIVPle subtypes among lion populations, refute the hypothesis that African lions consist of a single panmictic population. Our results suggest that extant lion populations derive from several Pleistocene refugia in East and Southern Africa (~324,000–169,000 years ago), which expanded during the Late Pleistocene (~100,000 years ago) into Central and North Africa and into Asia. During the Pleistocene/Holocene transition (~14,000–7,000 years), another expansion occurred from southern refugia northwards towards East Africa, causing population interbreeding. In particular, lion and FIVPle variation affirms that the large, well-studied lion population occupying the greater Serengeti Ecosystem is derived from three distinct populations that admixed recently.

Author Summary

The lion Panthera leo, a formidable carnivore with a complex cooperative social system, has fascinated humanity since pre-historical times, inspiring hundreds of religious and cultural allusions. Here, we use a comprehensive sample of 357 individuals from most of the major lion populations in Africa and Asia. We assayed appropriately informative autosomal, Y-chromosome, and mitochondrial genetic markers, and assessed the prevalence and genetic variation of the lion-specific feline immunodeficiency virus (FIVPle), a lentivirus analogous to human immunodeficiency virus (HIV) that causes AIDS-like immunodeficiency disease in domestic cats. We compare the large multigenic dataset from lions with patterns of genetic variation of the FIVPle to characterize the population-genomic legacy of lions. We refute the hypothesis that African lions consist of a single panmictic population, highlighting the importance of preserving populations in decline rather than prioritizing larger-scale conservation efforts. Interestingly, lion and FIVPle variation revealed evidence of unsuspected genetic diversity even in the well-studied lion population of the Serengeti Ecosystem, which consists of recently admixed animals derived from three distinct genetic groups.


Lion fossils trace to the Late Pliocene in Eastern Africa and the Early Pleistocene in Eastern and Southern Africa coincident with the flourishing of grasslands ~2–1.5 million years ago [1],[2]. By Mid Pleistocene (~500,000 years ago), lions occupied Europe and by the Late Pleistocene (~130,000–10,000 years ago) lions had the greatest intercontinental distribution for a large land mammal (excluding man), ranging from Africa into Eurasia and the Americas [3]. Lions were extirpated from Europe 2,000 years ago and within the last 150 years from the Middle East and North Africa. Today, there are less than 50,000 free-ranging lions [4] that occur only in sub-Saharan Africa and the Gir Forest, India (Figure 1A).
Figure 1. Geographic location of the lion samples and the variability of host and viral genetic markers among lion populations.
(A) Historical and current geographic distribution of lion, Panthera leo. A three-letter code pointing to a white dotted circle represents the geographic location of the 11 lion populations determined by Bayesian analyses [22] and factorial correspondence analyses [23] of the genetic distinctiveness of 357 lion samples (see text): GIR, Gir Forest, India; UGA, Uganda (Queen Elizabeth National Park); KEN, Kenya (Laikipia), SER, Serengeti National Park, Tanzania; NGC, Ngorongoro Crater, Tanzania; KRU, Kruger National Park, South Africa; BOT-I, southern Botswana and Kalahari, South Africa; BOT-II, northern Botswana; and NAM, Namibia. Green squares represent captive individual samples to explore the relationship of lions from more isolated/endangered/depleted areas: ATL, Morocco Atlas lions (n = 4); ANG, Angola (n = 2); and ZBW, Zimbabwe (n = 1). Deduced historical expansions (M1 and M2) are represented by red arrows (see text). (B) Haplotype frequencies observed in the 11 lion populations for nDNA (ADA and TF), and mtDNA (12S16S) genes, paralleled with the FIVPle serum-prevalence frequencies (black – sero-positive; gray – indeterminate; white – sero-negative). Population sample sizes are indicated within parenthesis. (C) Statistical parsimony networks of lion ADA, TF, and 12S16S haplotypes. Circle size is proportional to the haplotype frequency and crossbars represent the number of step mutations connecting haplotypes. The mtDNA haplotypes H5 and H6 are shaded gray as they were detected only in the individual samples from ANG, ATL, and ZBW, which do not group in unique population clusters (see text).
Understanding the broader aspects of lion evolutionary history has been hindered by a lack of comprehensive sampling and appropriately informative genetic markers [5][9], which in species of modern felids requires large, multigenic data sets due to its generally rapid and very recent speciation [10],[11]. Nevertheless, the unique social ecology of lions [12][14] and the fact that lions have experienced well-documented infectious disease outbreaks, including canine distemper virus, feline parvovirus, calicivirus, coronavirus, and lion feline immunodeficiency virus (FIVPle) [15][18] provide a good opportunity to study lion evolutionary history using both host and virus genetic information. Indeed, population genetics of transmitted pathogens can accurately reflect the demographic history of their hosts [19],[20]. Unlike other of the 36 cat species, lions have a cooperative social system (prides of 2–18 adult females and 1–9 males) and their populations can have high frequencies of FIVPle, a lentivirus analogous to human immunodeficiency virus (HIV), which causes AIDS-like immunodeficiency disease in domestic cats. FIVPle is a retrovirus that integrates into the host genome and is transmitted by cell-to-cell contact, which in felids occurs during mating, fighting and mother-to-offspring interactions. Thus, viral dissemination is a function in part of the frequency of contact between infected and naïve lions within and among populations. The virus is quite genetically diverse in lions [15],[18], offering a unique marker for assessing ongoing lion demographic processes.
To unravel lion population demographic history we used a large multigenic dataset. Distinct sets of markers may not necessarily yield similar inferences of population history, as coalescent times vary as a function of their pattern of inheritance [21]. There is also a large variance in coalescent times across loci sharing a common pattern of inheritance especially in complex demographical histories (Table 1). However, the accurate interpretation of the differences among loci can provide a more resolved and coherent population history, affording more-nuanced insights on past demographic processes, levels of admixture, taxonomic issues, and on the most appropriate steps for effective conservation and management of remaining populations.
Table 1. Expected and observed coalescent times for the different markers studied in lions according to their pattern of inheritance.
The goal of this study was to assess the evolutionary history of lion by (1) characterizing lion population structure relative to patterns of FIVPle genetic variation, (2) detect signatures of migration using both host and viral population genomics, and (3) reconstruct lion demographic history and discuss its implication for lion conservation. We assess genetic variation from 357 lions from most of its current distribution, including mitochondrial (mtDNA; 12S16S, 1,882 bp), nuclear (nDNA) Y-chromosome (SRY, 1,322 bp) and autosomal (ADA, 427 bp; TF, 169 bp) sequences, and 22 microsatellites markers. We further document patterns of FIVPle variation in lions (FIVPle pol-RT gene, up to 520 bp).


Population Structure of Lion

Genetic analyses of 357 lions from throughout the extant species range showed that paternally inherited nDNA (SRY) and maternal inherited (mtDNA) sequence variation was generally low (only one paternal SRY-haplotype and 12 mtDNA haplotypes; π = 0.0066) (Figure 1; Figure S1; Tables S1 and S2). The most common mtDNA haplotype H11 was ubiquitous in Uganda/Tanzania and parts of Botswana/South Africa, H1 was common in Southern Africa, and H7 and H8 were unique to Asian lions. The autosomal nDNA sequences showed fairly distinct patterns of variation (Figure 1; Figure S1). Of the five ADA haplotypes, A2 was the most common and most-widely distributed. The other four haplotypes, which are derived and much less common, included one (A5) that was fixed in Asian lions. The three TF haplotypes were more widely and evenly distributed.
Levels of population subdivision among lions were assessed using microsatellite and sequencing data. Eleven groups were identified using Bayesian analyses [22] and three-dimensional factorial correspondence analyses [23] (Figure 2; Table S3). Most clusters represented geographically circumscribed populations: Namibia (Nam), Kruger National Park (Kru), Ngorongoro Crater (Ngc), Kenya (Ken), Uganda (Uga), and Gir (Gir). Two distinct clusters were found in Botswana, Bot-I that included lions from southern Botswana and Kalahari (South Africa) (Fk = 0.24) and Bot-II found exclusively in northern Botswana (Fk = 0.18). Surprisingly, three distinct clusters were found in a single geographical locale (approximately 60×40 km square) in the large panmyctic population of the Serengeti National Park (Ser-I/Ser-II/Ser-III) (Fk = 0.18, 0.21, and 0.15, respectively).
Figure 2. Population structure analyses in lions.
(A) Bayesian population assignment test [22] of the 357 lions using 24 nDNA loci (ADA, TF, and 22 microsatellites) and mtDNA data, and considering K = 11 (11 populations). (B) Three-dimensional factorial correspondence analysis [23] (FCA) based on the 24 nDNA loci genotypes in the 357 lions. Axe 1, 2, and 3 represent 49.90% of the genetic variation observed. (C) FCA representation excluding the GIR lions. Axe 1, 2, and 3 represent 51.35% of the genetic variation observed. (D) FCA representation considering only the SER lions supportive of a three distinct population clusters subdivision (SER-I, SER-II, and SER-III).
Two captive lions from Angola (Ang), one from Zimbabwe (Zbw) and four Morocco Zoo Atlas lions (Atl; presently extinct from the wild) (Figure 1A) were included in the analyses to explore the relationship of lions from more isolated, endangered, or depleted areas. Ang and Zbw lions were assigned to Bot-II (q = 0.90 and 0.87; 90%CI: 0.47–1.00) and Kru (q = 0.85; 90%CI: 0.52–1.00) (Bayesian analyses [22]) populations, respectively, as expected based on their geographical proximity. However, these lions differed from Bot-II and Kru by up to 8 mtDNA mutations, sharing haplotypes with the Atl lions (H5 in Ang and H6 in Zbw) (Figure 1B and 1C). The Atl lions did not group in a unique cluster.
Both nDNA and mtDNA pairwise genetic distances among the 11 lion populations showed a significant relationship with geographic distance (R2 = 0.75; Mantel's test, P = 0.0097; and R2 = 0.15; Mantel's test, P = 0.0369; respectively) (Figure 3). The significant positive and monotonic correlation across all the scatterplot pairwise comparisons for the nDNA markers (bi-parental) was consistent with isolation-by-distance across the sampled region. However, the correlation between nDNA FST and geographic distance considerably decreased when the Asian Gir population was removed (R2 = 0.19; Mantel's test, P = 0.0065) suggesting that caution should be taken in interpreting the pattern of isolation-by-distance in lions. We further compared linearized FST estimates [24] plotted both against the geographic distance (model assuming habitat to be arrayed in an infinite one-dimensional lattice) and the log geographic distance (model assuming an infinite two-dimensional lattice). The broad distribution of lions might suggest a priori that a two-dimensional isolation-by-distance model would provide the best fit for the nDNA data (R2 = 0.25; Mantel's test, P = 0.0022), but instead the one-dimensional isolation-by-distance model performed better (R2 = 0.71; Mantel's test, P = 0.0476) (Figure S2).
Figure 3. Genetic differentiation of host and viral genetic markers with geographic distance.
Regression of lion pairwise FST (nDNA and mtDNA) and FIVPle (pol-RT) on geographic distance.
The pattern observed for the mtDNA (maternal) was more complex. While there was a significant relationship between mtDNA FST and geographic distance, there was an inconsistent pattern across broader geographic distances (Figure 3). This is partly due to the fixation or near fixation of haplotype H11 in six populations and the fixation of a very divergent haplotype H4 in Ken population (Figure 1B and 1C). The removal of the Ken population considerably increased the correlation between mtDNA FST and geographic distance (R2 = 0.27; Mantel's test, P = 0.0035). Thus, the null hypothesis of regional equilibrium for mtDNA across the entire sampled region is rejected despite the possibility that isolation-by-distance may occur regionally.
These contrasting nDNA and mtDNA results may be indicative of differences in dispersal patterns between males and females, which would be consistent with evidence that females are more phylopatric than males. Alternatively, selection for matrilineally transmitted traits upon which neutral mtDNA alleles hitchhike is possible, given the low values of nucleotide diversity of the mtDNA (π = 0.0066). A similar process has been suggested in whales (π = 0.0007) [25] and African savannah elephants (π = 0.0200) [26], where both species have female phylopatry and like lions, a matriarchal social structure. However, genetic drift tends to overwhelm selection in small isolated populations, predominantly affecting haploid elements due to its lower effective population size (Table 1). Therefore, we suggest that the contrasting results obtained for nDNA and mtDNA are more likely further evidence that lion populations underwent severe bottlenecks. The highly structured lion matrilines comprise four monophyletic mtDNA haplo-groups (Figure 4A; Figure S3). Lineage I consisted of a divergent haplotype H4 from Ken, lineage II was observed in most Southern Africa populations, lineage III was widely distributed from Central and Northern Africa to Asia, and lineage IV occurred in Southern and Eastern Africa.
Figure 4. Evolutionary relationships of the host and viral genetic markers among lion populations.
(A) Unrooted neighbour-joining (NJ) tree from nDNA genotypes of 24 loci (ADA, TF, and 22 microsatellites) in the 11 lion populations (left), and rooted NJ tree for the distinct mtDNA (12S16S, 1,882 bp) haplotypes in lion (right). The distinct mtDNA lineages were labelled I to IV. Bootstrap support (BPS) values >50 are indicated. (B) NJ tree of the 301 bp FIVPle pol-RT sequences. The distinct FIVPle subtypes were labelled A to F. BPS values are placed at each branchpoint and in parenthesis are the BPS values obtained for a tree established with 520 bp of FIVPle pol-RT sequence for a representative subset of individuals. (C) Distinctiveness of host and viral molecular genetics in lion populations.

Population Structure of FIVPle

Seroprevalence studies indicate that FIVPle is endemic in eight of the 11 populations but absent from the Asian Gir lions in India and in Namibia and southern Botswana/Kalahari regions (Nam/Bot-I) in Southwest Africa (Figure 1B). Phylogenetic analysis of the conserved pol-RT region in 117 FIV-infected lions depicted monophyletic lineages [15],[18] that affirm six distinct subtypes (A–F) that are distributed geographically in three distinct patterns (Figure 4B; Figure S4). First, multiple subtypes may circulate within the same population as exemplified by subtypes A, B and C all ubiquitous within the three Serengeti lion populations (Ser-I, Ser-II and Ser-III) and subtypes A and E within lions of Botswana (Bot-II) (Figure 4B and 4C and Figure S4). Second, unique FIVPle subtypes may be restricted to one location as subtype F in Kenya, subtype E in Botswana (Bot-II), subtype C in Serengeti, and subtype D in Krugar Park (Figure 4B and 4C and Figure S4). Third, intra-subtype strains cluster geographically, as shown by distinct clades within subtype A that were restricted to lions within Krugar Park, Botswana and Serengeti and within subtype B that corresponded to Uganda, Serengeti and Ngorongoro Crater lions (Figure 4B and 4C and Figure S4).
Not unexpectedly, FIVPle pairwise genetic distances, represented as population FST among the eight lion FIV-infected populations, were not significantly correlated with geographic distance (R2 = 0.08; Mantel's test, P = 0.165) (Figure 3), and affirms that patterns of viral dissemination do not conform to a strict isolation-by-distance model. Rather, the two distinct clusters observed (Figure 3) reflect the complex distribution of FIVPle among African lions. Indeed, despite the low geographic distance within East-African lion populations, the FIVPle genetic divergence showed a broader range in FST (0.03 to 0.79 for most of first cluster; Figure 3). By contrast, approximately half of the range in FST (0.26 to 0.69 for the second cluster; Figure 3) was observed among East and Southern Africa in spite of its large geographic separation. In contrast with the patterns observed in lions, linearized FST estimates [24] for FIVPle were better correlated with log geographic distance (two-dimensional lattice model) (R2 = 0.15) than with geographic distance (one-dimensional model) (R2 = 0.02), although in both cases the Mantel's test was not significant (P>0.2474) (Figure S2).

Natural History of Lions as Inferred from Lion and FIVPle Markers

The mtDNA coalescence dating suggested that the East African lineage I (Ken haplotype H4) had an old origin of ~324,000 years (95% CI: 145,000–502,000). Extant East African populations (Ken/Ngc/Ser-I/Ser-II/Ser-III) also showed a slightly significant higher nDNA allelic richness and genetic diversity (Table S4) relative to populations to the south (Kru/Nam/Bot-I/Bot-II) and north (Uga/Gir) (A = 2.43, 2.39, and 1.62, P = 0.021; HO = 0.64, 0.62, and 0.34, P = 0.019; respectively). Moreover, the FIVPle subtype diversity was higher in East African clades (exhibiting four out of the six known viral-strains), including the most divergent FIVPle subtype C (Figure 4B and 4C). These genetic data from lions and FIVPle is consistent with the older origin of extant East African lions, which is further supported by the oldest lion fossils discovered in East Africa [1].
Relative to East Africa, Southern lions have a slightly more recent mtDNA coalescence. Lineage II, found in Nam, Bot-II and Kru has an estimated coalescence of 169,000 years (95% CI: 34,000–304,000) and the more widespread lineage IV found in the Southern populations of Bot-I, Bot-II and Kru as well as the Eastern populations of Ser (I, II, and III), Ngc and Uga, coalesces ~101,000 years ago (95% CI: 11,000–191,000). However, the similar levels of nDNA genetic diversity, the occurrence of an exclusively Southern mtDNA lineage II and highly divergent FIVPle subtypes, FIVPle subtype D found only in Kru and subtype E exclusive to Bot-II, suggests that both East and Southern Africa were important refugia for lions during the Pleistocene. Therefore, the co-occurrence of divergent mtDNA haplotypes (6 to 10 mutations; Figure 1B and 1C) in southern populations may be the consequence of further isolation within refugia during colder climatic periods. Contemporary fragmentation of lion populations could further explain the results of nested-clade phylogeographical analysis (NCPA [27]) (Figure S5), which inferred restricted gene flow with isolation-by-distance between mtDNA haplotypes H9 (Bot-II) and H10 (Kru) (χ2 = 10.00, P = 0.0200), between haplotypes H1 (Bot-II/Nam) and H2 (Kru) (χ2 = 71.00, P≤0.0001), and between haplotypes H9–H10 (Bot-II/Kru) and haplotypes H11–H12 (Bot-I/Kru/Ser/Ngc/Uga) (χ2 = 187.83, P≤0.0001).
Further isolation within refugia (sub-refugia) may also have occurred in East Africa. This is suggested by the distinctive mtDNA haplotype H4 and the unique FIVPle subtype F found in the Kenya population, which may have resulted from reduced gene flow across the Rift valley, a scenario that has been suggested for several bovid and carnivore populations (see [28] and references therein).
The best example of concordance between host genome markers and viral transmission patterns is observed in the Serengeti National Park in Tanzania. Our previous findings described markedly high levels of FIVPle subtype A, B and C circulating within the Serengeti lion population to such an extent that 43% of the lions sampled were multiply-infected with two or three subtypes [15],[18] and were hypothesized to represent recent admixture of three formerly separated populations. Such result is confirmed here by lion genomic markers (Figure 2). Further, although lions within the Serengeti can be assigned to one of three populations (Ser-I, Ser-II or Ser-III) by host genomic markers, FIVPle subtypes are distributed ubiquitously in all three, characteristic of rapid horizontal retroviral transmission subsequent to host population admixture. The possible isolating mechanism remains to be elucidated as there is no apparent barrier to gene flow in this ecosystem.

Genomic Signatures Left by Migration

Based on patterns of genetic diversity and phylogenetic analysis of lion nDNA/mtDNA and FIVPle markers, we propose a scenario of a period of refugia/isolation in the Late Pleistocene followed by two major lion expansions across Africa and Asia. The first expansion, supported by the mtDNA NCPA [27]2 = 690.00, P≤0.0001; Figure S5), was a long-distance colonization of mtDNA lineage-III (Gir/Atl/Ang/Zbw) around 118,000 years ago (95% CI: 28,000–208,000), with subsequent fragmentation of haplotypes H5–H6 into Central and North Africa and haplotypes H7–H8 into West Asia (M1- Figure 1A). Support for this initial expansion is also found in nDNA. The ADA haplotype A5 fixed in Gir in also present in Ken, Ser-II, and Ser-III, suggesting that lions likely colonized West Asia from the East Africa refugia (Figure 1B). Such an expansion may have been favored by the start of a warmer and less arid period in Africa 130,000–70,000 years ago [29]. This “out-of-Africa event” would have occurred much later than the initial lion expansion through Eurasia based on fossils (~500,000 years ago) [3]. It is likely that multiple lion expansions occurred in the Pleistocene, as occurred with humans [21].
A second, more recent lion expansion probably occurred at the Pleistocene/Holocene transition, this one from Southern Africa toward East Africa (M2- Figure 1A, Figure 3). This is reflected in the mtDNA linage IV, where haplotypes present in Southern lions are basal (older) to those found in the East. Overall, mtDNA population nucleotide diversity decreases from Southern to East Africa (Figure 1B and 1C), a finding supported by pairwise mismatch analysis [30] (raggedness, r = 0.086; P<0.001). The fixation of mtDNA haplotype H11 in Bot-I (otherwise fixed only in East Africa populations) suggests that the colonizing lions expanded northwards from the Kalahari Desert, which included bush, woodland and savannah habitats during the climatic fluctuations of the Pleistocene [31]. This expansion would have occurred relatively recently as the single rare tip mtDNA haplotype H12, found only in Ser-I, is derived from the interior widespread haplotype H11 (~14,000–7,000 years; given one mtDNA substitution every 7,000 years; Table 1). This expansion is also supported by FIVPle subtype A where haplotypes present in Southern lions (Kru and Bot-I) are basal to those found in the East (Ser-I, Ser-II and Ser-III) and a decrease of nucleotide-diversity of this FIVPle subtype is observed from Southern (π = 0.15) to Eastern Africa (π = 0.03) (Figure 3B). Interestingly, a similar northward colonization process from Southern Africa has been suggested for some of the lion preys, namely the impala, greater kudu, and wildebeest [32],[33].

Utility of Population Genomic Datasets

If we had restricted our inferences to mtDNA, we might have concluded that East African lion populations, which are fixed or nearly fixed for haplotype H11, went extinct during the Pleistocene/Holocene transition (similar to the well known mega-fauna extinctions of the Late Pleistocene [34]) and were then colonized by Southern populations. However, our population genomics data better fit a scenario of lion population expansion and interbreeding rather than simple replacement. First, genetic diversity and allelic richness at nDNA are slightly higher in East Africa populations relatively to those in Southern Africa. This is contrary to the expected pattern of population expansion in which there is usually a progressive decline in genetic diversity and allelic richness. Second, Ser lions carry two diverse FIVPle subtypes found only in East Africa (B–C), and not only FIVPle subtype A, which was presumably introduced in East Africa coincidently with the mtDNA expansion event northwards from South. Third, the East African FIVPle subtype B found in Uga/Ser-I/Ser-II/Ser-III/Ngc showed evidence of a population expansion (raggedness, r = 0.004; P<0.01; Fs = −20.37; P<0.00001) and the highest nucleotide diversity observed within FIVPle subtypes (π = 0.09). Four, the FIVPle subtype diversity is higher in East African clades (four out of the six viral strains).
The utility of FIVPle pol-RT as a marker of lion population structure and natural history is that it can be informative on a contemporaneous time scale, though it may be less useful at capturing more ancient demographic events. The extreme divergence among FIVPle subtypes, considered with high sero-prevalence in eight of the 11 lion populations, and combined with patterns of geographic concordance, support the hypothesis that FIVPle is not a recent emergence within modern lions [35]. Populations that harbor one private FIVPle subtype such Ken (subtype F), Bot-II (subtype E), and Kru (subtype D) must have been sufficiently isolated for enough time for the virus to evolve into unique subtypes, a result corroborated by the high nDNA and mtDNA genetic structure present in these lion populations (Figure 4). Thus, it is possible that the initial emergence of FIVPle pre-dates the Late-Pleistocene expansions of contemporary lion populations [36], but present day distributions are more useful indicators of very recent host population dynamics, a result also observed with FIVPco in a panmictic population of pumas in western North America [19].

Conservation Implications

Accurate interpretation of past and contemporary population demographic scenarios is a primary goal for the effective conservation of endangered species. In this study, we found substantial population subdivision, reduced gene flow, and large differences in FIVPle sequence and sero-prevalence among lion populations, as well as evidence of historic secondary contact between populations (Figure 3C; Table S4 to S9). The very low population level of mtDNA nucleotide diversity, the number of haplotypes private to a single population (Figure 1), and probably also the lack of SRY genetic variation across all male lions (haplotype S1, n = 183) suggests that lion numbers diminished considerably following the Late Pleistocene. The last century reduction in lion distribution further eroded its genomic diversity, and microsatellite variation suggested recent population bottlenecks in seven out of the 11 populations (standardized differences test, P<0.05; Table S5) [37].
Although we did not explicitly try to address the adequacy of lion subspecies designations (currently only one African subspecies is widely recognized) [38],[39], we provided strong evidence that there is no evidence of substantial genetic exchange of matrilines among existing populations as the AMOVA [40] within-population component was uniformly high in all distinct subdivision scenarios (ΦST≈0.920; P<0.0001; three-six groups; Table S6). Similarly, significant population structure was detected from nDNA (FST = 0.18), with low levels of admixture evident from Bayesian analysis [22] (α = 0.033). Therefore, employing a bottom-up perspective that prioritizes populations, rather than large-scale units (e.g. all African lions), might preserve and maintain lion diversity and evolutionary processes most efficiently [41].

Material and Methods

Study Site, Sampling, and Molecular Genetic Analyses of Lions

A total of 357 individuals were obtained across most of the lion range in Africa and Asia (Figure 1A; Table S1). Genetic variation among lion specimens was assessed using maternal (12S and 16S genes), paternal (SRY gene) and bi-parental autosomal (22 microsatelite loci, and the ADA and TF genes) markers (GenBank accession numbers: FJ151632–FJ151652). Analyses of mtDNA in Panthera species are complicated by the presence of a 12.5 kb mtDNA integration into chromosome F2 [42]. Accordingly, mtDNA specific primers were designed for the 12S and 16S genes (Table S2) and we used long-range PCR amplification. We designed primers to amplify segments of the ADA (exon 10 and intron 10) and the TF (intron 3) genes (Table S2), two of the most variable protein loci in lion populations [5], localized on the domestic cat Felis catus chromosome A3p and C2q, respectively. The Y-chromosome SRY-3′UTR gene was also amplified [43].
PCR products were amplified from 50 ng of genomic DNA in a 25 µL reaction system containing 1.5 mM MgCl2, 1.0 mM dNTPs, 0.25 units of AmpliTaq Gold DNA polymerase (Applied Biosystems), and 1× PCR buffer II; the amplification protocol was: denaturation 10 min at 95°C, a touch-down cycle of 95°C for 30 s, 52°C for 60 s decreased by 1°C in the next cycle for 10 cycles, 72°C for 120 s, then 35 amplification cycles of 95°C for 30 s, 52°C for 60 s, and 72°C for 120 s, followed by an extension of 10 min at 72°C. PCR products were sequenced on an ABI 377. Sequences were aligned and cleaned using SEQUENCHER (Gene Codes).
Twenty two polymorphic microsatellite loci (20 dinucleotide repeats: FCA006, FCA008, FCA014, FCA069, FCA077, FCA085, FCA091, FCA098, FCA105, FCA126, FCA129, FCA139, FCA205, FCA208, FCA211, FCA224, FCA229, FCA230, FCA247, and FCA281; and two tetranucleotide repeats: FCA391 and FCA441) were amplified [44]. Microsatellites were scored in an ABI 377 and analyzed using Genescan 2.1 and Genotyper 2.5. These loci are located on 11 of the 19 F. catus chromosomes, occurring in different linkage groups or at least 12 centimorgans apart [44],[45].

Sero-Prevalence and Molecular Genetics of FIVPle

Western blots using domestic cat and lion FIV as antigen were performed as previously described [46],[47]. The supernatant from virus-infected cells was centrifuged at 200 g for 10 min at 5°C. The resultant supernatant was centrifuged at 150,000 g at 4°C for 2 hours. Pelleted viral proteins were resuspended in 1/20th of the original volume and total protein content was assayed using the Biorad Protein Assay. Twenty mg of viral protein were run on 4–20% Tris-Glycine gels and transferred to PDVF membranes (BioRad). Membrane strips were exposed 2–12 h to a 1:25 or 1:200 dilution of serum or plasma. After washing, samples were labeled with goat anti-cat HRP or phosphate conjugated antibody (KPL laboratories) at a 1:2000 dilution, washed, and incubated in ECL Western Blotting detection reagents (Amersham Biosciences) for 2 min, then exposed to Lumi-Film Chemiluminescent Detection Film (Boehringer Mannheim) or incubated in BCIP/NBP phosphatase substrate (KPL laboratories) for 15 min [46][48]. Results were visualized and scored manually based on the presence and intensity of antibody binding to the p24 gag capsid protein.
Nested PCR amplification of partial FIVPle pol-RT was performed [18],[46]. Briefly, first round PCR reactions used 100 ng of genomic DNA, 2.5 mM MgCl2 and an annealing temperature of 52°C. Second round PCR reactions used identical conditions with 1–5 µl of first-round product as template. All PCR products were sequenced as described above for lion genetic analyses (GenBank accession numbers: AY549217–AY552683; AY878208–AY878235; FJ225347–FJ225382).

Statistical Analyses

We used the Genetix 4.02 [49], Genepop 3.3 [50], Microsat [51], and DnaSP 4.10 [52] to calculate the following descriptive statistics: (i) percentage of polymorphic loci (P95), number of alleles per locus (A), observed and expected heterozygosity (HE and HO), and number of unique alleles (AU); (ii) assess deviations from HWE; (iii) estimate the coefficient of differentiation (FST), and (iv) nucleotide (π) and haplotype (h) diversity.
We tested the hypothesis that all loci are evolving under neutrality for both the lion and the FIVPle data. For frequency data, we used the method described by Beaumont and Nichols [53] and implemented in Fdist (​.html). The FST values estimated from microsatellite loci plotted against heterozygosity showed that all values fall within the expected 95% confidence limit and consequently no outlier locus were identified. For sequence data (lion nDNA/mtDNA and FIVPle pol-RT), we ruled out any significant evidence for genetic hitchhiking and background selection by assessing Fu and Li's D* and F* tests [54] and Fu's FS statistics [55].
A Bayesian clustering method implemented in the program Structure [22] was used to infer number of populations and assign individual lions to populations based on multilocus genotype (microsatellites) and sequence data (ADA, TF, and mtDNA genes) and without incorporating sample origin. For haploid mtDNA data, each observed haplotype was coded with a unique integer (e.g. 100, 110) for the first allele and missing data for the second (Structure [22] analyses with or without the mtDNA data were essentially identical). For K population clusters, the program estimates the probability of the data, Pr(X|K), and the probability of individual membership in each cluster using a Markov chain Monte Carlo method under the assumption of Hardy-Weinberg equilibrium (HWE) within each cluster. Initial testing of the HWE in each of the populations defined by the geographic origin of sampling revealed no significant deviation from HW expectations with the exception of Ser and Bot population (later subdivided by Structure [22] in 3 and 2 clusters, respectively; such deviations from HW expectations were interpreted as evidence of further population structuring). We conducted six independent runs with K = 1–20 to guide an empirical estimate of the number of identifiable populations, assuming an admixture model with correlated allele frequencies and with burn-in and replication values set at 30,000 and 106, respectively. Structure also estimates allele frequencies of a hypothetical ancestral population and an alpha value that measures admixed individuals in the data set. The assignment of admixed individuals to populations using Structure [22] has been considered in subsequent population analyses. For each population cluster k, the program estimates Fk, a quantity analogous to Wright's FST, but describing the degree of genetic differentiation of population k from the ancestral population.
Patterns of gene flow and divergence among populations were described using a variety of tests. First, to visualize subtle relationships among individual autosomal genotypes, three-dimensional factorial correspondence analyses [23] (FCA) were performed in Genetix [49], which graphically projects the individuals on the factor space defined by the similarity of their allelic states. Second, neighbor-joining (NJ) analyses implementing the Cavalli-Sforza & Edwards' chord genetic distance [56] (DCE) were estimated in Phylip 3.6 [57], and the tree topology support was assessed by 100 bootstraps. Third, the difference in average HO and A was compared among population groups using a two-sided test in Fstat [58], which allows to assess the significance of the statistic OSx using 1,000 randomizations. Four, the equilibrium between drift and gene flow was tested using a regression of pairwise FST on geographic distance matrix among all populations for host nDNA(microsatellites)/mtDNA and FIVPle data. A Mantel test [59] was used to estimate the 95% upper probability for each matrix correlation. Assuming a stepping stone model of migration where gene flow is more likely between adjacent populations, one can reject the null hypothesis that populations in a region are at equilibrium if (1) there is a non-significant association between genetic and geographic distances, and/or (2) a scatterplot of the genetic and geographic distances fails to reveal a positive and monotonic relationship over all distance values of a region [60]. We also evaluated linearized FST [i.e. FST/(1−FST)] [24] among populations. We tested two competing models of isolation-by-distance, one assuming the habitat to be arrayed in an infinite one-dimensional lattice and another assuming an infinite two-dimensional lattice. Both models showed that genetic differentiation increased with raw and log-transformed Euclidean distances, respectively [24]. We determined the confidence interval value of the slope of the regression for the nDNA data using a non parametric ABC bootstrap [61] in Genepop 4.0 [62].
The demographic history of populations was compared using a variety of estimators based on the coalescence theory. First, signatures of old demographic population expansion were investigated for mtDNA and FIVPle pol-RT haplotypes using pairwise mismatch distributions [63] in DnaSP [52]. The goodness-of-fit of the observed data to a simulated model of expansion was tested with the raggedness (r) index [64].
Second, the occurrence of recent bottlenecks was evaluated for microsatellite data using the method of Cornuet & Luikart [37] in Bottleneck [65] and using 10,000 iterations. This approach, which exploits the fact that rare alleles are generally lost first through genetic drift after reduction in population size, employs the standardize differences test, which is the most appropriate and powerful when using 20 or more polymorphic loci [37]. Tests were carried out using the stepwise mutation model (SMM), which is a conservative mutation model for the detection of bottleneck signatures with microsatellites [66].
Third, to discriminate between recurrent gene flow and historical events we used the nested-clade phylogeographical analysis [27],[67] (NCPA) for the mtDNA data. When the null-hypothesis of no correlation between genealogy and geography is rejected, biological inferences are drawn using a priori criteria. The NCPA started with the estimation of a 95% statistical parsimony [68] mtDNA network in Tcs 1.20 [69]. Tree ambiguities were further resolved using a coalescence criteria [70]. The network was converted into a series of nested branches (clades) [71],[72], which were then tested against their geographical locations through a permutational contingency analysis in GeoDis 2.2 [73]. The inferences obtained were also corroborated with the automated implementation of the NCPA in ANeCA [74]. To address potential weaknesses in some aspects of the NCPA analysis [75],[76], we further validated the NCPA inferences with independent methods for detecting restricted gene-flow/isolation-by-distance (using matrix correlation of pairwise FST and geographic distance) and population expansion (using pairwise mismatch distributions).
Four, to test the significance of the total mtDNA genetic variance, we conducted hierarchical analyses of molecular variance [40] (AMOVA) using Arlequin 2.0 [77]. Total genetic variation was partitioned to compare the amount of difference among population groups, among populations within each groups, and within populations.
Phylogenetic relationships among mtDNA and FIVPle pol-RT sequences were assessed using Minimum evolution (ME), Maximum parsimony (MP), and Maximum likelihood (ML) approaches implemented in Paup [78]. The ME analysis for mtDNA consisted of NJ trees constructed from Kimura two-parameter distances followed by a branch-swapping procedure and for FIVPle data employed the same parameter estimates as were used in the ML analysis. The MP analysis was conducted using a heuristic search, with random additions of taxa and tree-bisection-reconnection branch swapping. The ML analysis was done after selecting the best evolutionary model fitting the data using Modeltest 3.7 [79]. Tree topologies reliability was assessed by 100 bootstraps. For the FIVPle data, the reliability of the tree topology was further assessed through additional analyses using 520 bp of FIVPle pol-RT sequences in a representative subset of individuals.
The time to the most recent common ancestor (TMRCA) for the ADA and TF haplotypes was estimated following Takahata et al. [80], where we calculate the ratio of the average nucleotide differences within the lion sample to one-half the average nucleotide difference between leopards (P. pardus) and lions and multiplying the ratio by an estimate of the divergence time between lions and leopards (2 million years based on undisputed lion fossils in Africa) [81],[82]. The mtDNA TMRCA was estimated with a linearized tree method in Lintree [83] and using the equation H = 2μT, where H was the branch height (correlated to the average pairwise distance among haplotypes), μ the substitution rate, and T the divergence time. Leopard and snow leopard (P. uncia) sequences were used as outgroups. Inference of the TMRCA for microsatellite loci followed Driscoll et al. [6] where the estimate of microsatellite variance in average allele repeat-size was used as a surrogate for evolutionary time based on the rate of allele range reconstitution subsequent to a severe founder effect. Microsatellite allele variance has been shown to be a reliable estimator for microsatellite evolution and demographic inference in felid species [6].

Supporting Information

Genetic variation of 12S–16S (mtDNA) and ADA and TF (nDNA) genes in lions. (A) Haplotypes and variable sites for the 12S–16S mtDNA region surveyed in lions (total length 1,882 bp). Position 1 corresponds to position 1441 of the domestic cat (Felis catus) mtDNA genome (GenBank U20753). The “-” represents a gap and “.” matches the nucleotide in the first sequence. Shading indicates a fixed difference in the mtDNA lineage. (B) Haplotypes and variable sites for the ADA gene segment surveyed in lions (total length 427 bp). (C) Haplotypes and variable sites for the TF gene segment surveyed in lions (total length 427 bp).
Genetic variation of 12S–16S (mtDNA) and ADA and TF (nDNA) genes in lions. (A) Haplotypes and variable sites for the 12S–16S mtDNA region surveyed in lions (total length 1,882 bp). Position 1 corresponds to position 1441 of the domestic cat (Felis catus) mtDNA genome (GenBank U20753). The “-” represents a gap and “.” matches the nucleotide in the first sequence. Shading indicates a fixed difference in the mtDNA lineage. (B) Haplotypes and variable sites for the ADA gene segment surveyed in lions (total length 427 bp). (C) Haplotypes and variable sites for the TF gene segment surveyed in lions (total length 427 bp).
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Linearized genetic differentiation of host and viral genetic markers with geographic distance. Regression of linearized FST estimates [24] for lion (nDNA and mtDNA) and FIVPle (pol-RT) genetic data plotted both against the geographic distance (model assuming habitat to be arrayed in an infinite one-dimensional lattice; one-dimension isolation-by-distance [IBD]) and the log geographic distance (model assuming an infinite two-dimensional lattice; two-dimension isolation-by-distance) on geographic distance.
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Phylogenetic relationships of the 12S–16S mtDNA lion haplotypes. Neighbour-joining tree of the 1,882 bp 12S–16S mtDNA sequences. Bootstrap values are placed at each branchpoint for the minimum evolution/maximum parsimony/maximum likelihood analyses, respectively (ME/MP/ML). Outgroups: Ppa – leopard, Panthera pardus; Pun – snow-leopard, Panthera uncia. The symbol (•) represents nodes with bootstrap support <50 or an inferred polytomy in the bootstrap 50% majority-rule consensus tree.
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Phylogenetic relationships of the FIVPle pol-RT sequences. Neighbour-joining tree of the 301 bp FIVPle pol-RT sequences. The distinct FIVPle subtypes were labelled A to F. Bootstrap (BPS) values are placed at each branchpoint (ME/MP/ML) and in parenthesis are the BPS values obtained for a tree established with 520 bp of FIVPle pol-RT sequence for a representative subset of individuals.
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Nested design and summary results of the nested clade phylogeographic analysis (NCPA) for lion mtDNA data. (A) Nested design of the mtDNA haplotype network used for the NCPA. (B) Summary results of the NCPA. RGF/IBD - Restricted gene flow/isolation by distance. LDC/FR – long distance colonization/fragmentation.
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List of the lion samples used in this study.
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Primers used to amplify the mtDNA (12S–16S) and nDNA (ADA and TF) portions surveyed in this study.
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Structure cluster assignment results of 357 lions based on nDNA (ADA, TF, and 22 microsatellites) and mtDNA markers. Burn-in and replication values set at 30,000 and 1,000,000, respectively.
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Gene diversity and frequency values in lion populations.
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Bottleneck analysis in lion populations using the standardized differences test and the stepwise mutation model (SMM).
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Results of the hierarchical AMOVA in lions for four different geographical scenarios.
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Lion population pairwise FST estimates. Below the diagonal mtDNA data (12S–16S) and above the diagonal microsatellite data (22 loci).
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Taxon specific unique nDNA alleles in lion populations (FCA-microsatellites and ADA locus).
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Summary statistics for FIVPle data.
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Tissues were obtained in full compliance with specific Federal Fish and Wildlife Permits. The content of this publication does not necessarily reflect the views or policies of the Department of Health and Human Services, nor does mention of trade names, commercial products, or organizations imply endorsement by the U.S. Government. We would like to thank A. Beja-Pereira, M. Branco and J. Martenson for suggestions and technical assistance. Comments made by the Associate Editor A. Estoup and three anonymous referees improved a previous version of this manuscript.

Author Contributions

Conceived and designed the experiments: AA JLT SJO WEJ. Performed the experiments: AA JLT MER WEJ. Analyzed the data: AA JLT MER SJO WEJ. Contributed reagents/materials/analysis tools: AA JLT MER JPS CP CW HW GH LF PS LS MD PJF KAA KCP GM DW MB SJO WEJ. Wrote the paper: AA JLT JPS CP SJO WEJ.


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