People spend more than a third of their disposable income on rent across large parts of England, a BBC investigation has found.

Analysis shows the average rent of a one-bedroom property in almost half of all districts, boroughs and cities would cost more than 30% of the median take-home salary for the area.
The problem is most acute in London and the South East.
The government said it was committed to building more affordable homes.
BBC England's data unit analysed the average rents for different types of property in each local authority area and the median weekly wage for those areas, using figures published by the Office for National Statistics.
According to Shelter and the Joseph Rowntree Foundation, spending more than a third of your disposable income on rent or a mortgage means you may not be able to afford other basic needs.
Based on average rents and median weekly wages, our investigation found:
Dan Wilson Craw, a policy manager at Generation Rent, which campaigns for affordable housing, said: "Across London and the South East, the only option for average earners is to squeeze themselves into ever more crowded flat shares.
"This might work for some, but it's a completely unsustainable situation for anyone who wants to settle down. Unless rents start coming down, the capital and its hinterland will start losing workers and that will weaken the national economy."
Kerry Taylor works as a teaching assistant in a special school in Stratford, earning about £350 a week in take-home pay.
She pays £825 a month to rent a room in a shared house in Lambeth, south London, where she has her own bathroom and kitchen.
She said: "I would like to get on the housing ladder but I have resigned myself to the fact that working in social care isn't going to get you the right salary, let alone give you the chance to save up for anything.
"I would love to own a home so that no one can kick me out, as I now have to leave my house as the landlady wants to sell.
"I've lived in Manchester, Edinburgh and my home town in Yorkshire but I've never been happier and more comfortable than in London. This is really important to me and I have created a great network of friends so the idea of moving and starting again somewhere else, as I've done twice before, is too exhausting to think about.
"I hope it doesn't come to having to leave but I suppose I can't rule it out completely if it gets that desperate."
Freelance musician and composer Matt Winkworth has been renting houses around Oxford for 10 years, but to gain security he has decided to buy a narrow boat, which he intends to live in.
The 31-year-old said: "Buying a boat is something that I've been thinking about doing for a long time.
"I've thought about getting a mortgage, but I would have to move out of the area that I work [in], it's impossible to buy anywhere in the South East.
"[Living in the boat] will be the first time I have lived on my own."
Sophie Mogridge moved to Oxford last year and lives in a three-bedroom house with two friends.
The trainee teacher has resigned herself to the fact that she can no longer live in the city, which is considered the most unaffordable place to live in Britain.
She spends £515 a month on rent but, when bills are taken into account, she spends half her £18,000-a-year income on accommodation.
The 23-year-old said: "My rent is incredibly high but my salary isn't as high.
"That means I'm not able to do all the social things that all my friends [outside of Oxford] can do.
"I'm actually moving to Banbury so I can afford to enjoy life."
£550
30% of average monthly take home wage in England
£694
Average rent for one-bed flat
£867 Average rent for three-bed home
£760 Average rent for two-bed home
£641 Average rent for a studio flat
£382 Average rent for house share
Our figures showed 142 out of 324 areas where the average rent for a one bedroom property was higher than 30% of take-home wage.  None were in the north of the country except Manchester, Salford and York.
Copeland in Cumbria seems to have that magic mix of cheap rents and high wages.
However, the data does not tell the full story.  Copeland is home to the Sellafield nuclear fuel reprocessing and nuclear decommissioning site. There are also proposals for a new nuclear power station, the Moorside project, to the north of the current Sellafield site. The energy industry in Copeland also includes the Walney Wind Farm offshore.
Lynne Atkinson, of Cumbrian lettings agency Grisdales, said: "Nuclear power does skew the figures when it comes to wages as there are some very well-paid jobs in the area.
"It is good value for money renting in Copeland and the market is very strong. It is cheaper than other areas in the North West, such as the Lakes. There has also been a lot of buy to let investment."
Campbell Robb, chief executive of Shelter, said: "Our chronic housing shortage means private renting is no longer a stepping stone for people starting out in life - it's where a quarter of families have to live.
"And with sky high rents eating up a huge chunk of people's monthly income, it's sadly no surprise that at Shelter we're hearing from growing numbers of families who are struggling just to cover the cost of the basics, and keep a roof over their heads."
Explore the full dataset here
Henry Gregg, assistant director of communications and campaigns at the National Housing Federation said: "These figures provide yet more evidence of how seriously unaffordable renting is in this country. Sky-high rents mean unstable and uncertain living situations are becoming the norm."
Our investigation used the median average, which describes the middle value in a set.
The median monthly take home pay in England is about £1,833, after tax and national insurance contributions.
That means rent should take up no more than £550.  The England average price of a room in a house is £382, while a one-bedroom property costs £694.
Alan Ward, who chairs the Residential Landlords Association said there was a rental "supply crisis" because the government had focussed on home ownership.
According to a survey carried out by the association, 84% of landlords were considering raising rent due to "punitive tax changes by the government".
A Department of Communities and Local Government spokesman said: "More than 300,000 people have been helped in homeownership through government-backed schemes since 2010, while almost 900,000 more homes have been delivered since the end of 2009.
"But we know there is more to do. That's why we've doubled the housing budget, including investing £8bn in an extra 400,000 quality affordable homes to rent and buy.
"We're also extending shared ownership, giving more people the chance to buy a home with a deposit of as little as £1,500."
Reporting team: Peter Sherlock, Daniel Wainwright, Paul Bradshaw, Steven Connor


Publisher Mills & Boon is to launch a series of "Modern Girl's Guides" containing "vital advice every woman needs to survive the modern world".

Tongue-in-cheek guides to working and drinking will be published in November, with more titles to follow.
Written by two unidentified authors, they will be published under the pen name Ada Adverse.
The series is a departure from Mills & Boon's normal output of steamy romance novels.
According to the publisher, the mock guides will juxtapose black-and-white archive photos of women with "witty 21st Century commentary".
A sample page from one of the titles, on the subject of friends, tells of two girls named Linda and Simone "who are going to a party".
"Linda is going to keep an eye on her pedometer and tell Simone when they have danced enough to leave and get doner kebabs," the text reads.
The initiative comes in the wake of the success of the Ladybird Books for Grown-ups series, a spoof on the classic children's books, which has now achieved more than two million sales.
Mills & Boon executive Lisa Milton said the books would offer "affordable, fun escapism".
"We couldn't resist bringing together an inspirational author and fabulous archive photos to create the Mills & Book Modern Girl's Guides," she continued.
"Whether its work or sex, going out or staying in - Mills & Boon has the guide for you."
According to a press release, Ada Adverse is "the world's leading authority on Having Fun" and is the author of more than 100 books.
The Mills & Boon Modern Girl's Guide to Working 9 to 5 and The Mills & Boon Modern Girl's Guide to Happy Hour will be published on 3 November.
They will be followed by The Mills & Boon Modern Girl's Guide to Helping Yourself on 29 December.
A fourth book, The Mills & Boon Modern Girl's Guide to Happy Endings, will hit shelves on 26 January.
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The Liberal Democrats will vote against the Welsh government's budget plans for local councils, in protest at cuts for rural authorities.

Powys, Ceredigion and Monmouthshire all face cuts of more than 3%, while Cardiff will lose just 0.1%.
Welsh Lib Dem leader Kirsty Williams said: "There is absolutely no way we could support cuts that hit rural areas so badly."
The Welsh government said councils had agreed to the settlement.
Without an overall majority in the assembly, Labour ministers need support from at least one other party to get the budget passed.
The Lib Dems claimed a funding floor protected rural council budgets in 2015/16, but the measures had not been repeated for 2016/17.
"We are calling on the Welsh government to introduce a rural grant that would limit the reductions for the councils worst hit," said Ms Williams.
"In the longer term, we need a fairer funding formula that recognises the higher costs that rural authorities face in delivering key public services."
A Welsh government spokeswoman said the formula used to distribute the available funds "is agreed each year in partnership with local government".
She added that the funding floor was not applied for 2016/17 as the settlement was "better than expected" with an overall cut of 1.4% compared to 3.4% in 2015/16.


An HGV lorry driver who was caught driving the wrong way along a dual carriageway in Flintshire after drinking whisky has been jailed.

Nicolae Gorea, 40, from Moldova, was more than twice the drink drive limit when he drove on the A548 in the Deeside Industrial Park on Good Friday.
A fellow trucker tried to stop him by driving to a roundabout - but he drove off, Flintshire magistrates heard.
Gorea admitted dangerous driving, drink driving and fraud.
District Judge Gwyn Jones jailed Gorea for 42 weeks and banned him from driving for two years.
The court heard he was spotted driving the wrong way along the busy road at about 22:35 BST by another lorry driver, Brian O'Connor, who tried to flag him down before driving to a roundabout to try and stop him.
But, unable to understand him, the defendant drove on, the court heard.
Mr O'Connor called the police before driving to another roundabout and eventually making Gorea stop and get out of his vehicle.
When police arrived they noticed there was damage to the front of his lorry, indicating it had been in contact with street furniture.
Checks also showed that Gorea was wanted by The Met police following a fraudulent driving licence application when a foreign Romanian driving licence had been used.
Interviewed, he said he had parked up for the night at Deeside Industrial Park on 14 April, drank between 200 and 300 millilitres of whisky and then decided to move the vehicle.
He admitted he had driven the wrong way and said he "was tired, confused and drunk."
The defendant also received a notice to be deported.
"You were driving a Romanian registered heavy goods vehicle with a 40 foot trailer on one of the arterial roots into north Wales on a busy Easter weekend," the judge said.
"You were significantly under the influence of alcohol and the manner of your driving clearly showed that."
He also praised Mr O'Connor for trying to stop the defendant and said his actions may well have helped others not to be hurt.


Four people have escaped from a burning house after an arson attack in Londonderry.

12 April 2016 Last updated at 21:14 BST
Police have yet to establish a motive for the blaze and have appealed for information.
BBC News NI's Keiron Tourish reports.


A 64-year-old man has died after his car collided with a pick-up truck near Newport.

His Vauxhall Astra convertible collided with an Isuzu pick-up on the A467 between Bassaleg roundabout and Rogerstone.
The road was closed following the crash on Friday at about 16:30 BST.
The driver and passenger of the truck are being treated at the city's Royal Gwent Hospital.
It is understood their injuries are not life threatening.
Gwent Police are appealing for witnesses.


Almost 1,000 people have signed an online petition to stop a hotel chain building in the UK's smallest city.

A 63-bedroom Premier Inn is included in plans to develop land at Glasfryn Road in St Davids, Pembrokeshire.
St Davids is renowned for its cathedral, independent shops and businesses, and is bidding to become the UK's 2021 City of Culture.
A spokesman for Premier Inn said issues raised by the petition would be clarified at public exhibitions.
The petition claims the hotel would "undermine" local businesses and "pave the way for more big chains" to come to the city.
The Glasfryn Road development includes 75 new homes, 40 of which would be available to rent from Pembrokeshire Housing Association and reserved for people who live locally.
It is hoped homes sold on the open market will help fund a new swimming pool to replace the one which closed in 2007.
The petition, set up on behalf of the No Premier Inn St Davids (NOPI) group, said the housing development "does not need and should not support a Premier Inn".
It said the hotel was part of a "large multi-national company" and "could potentially pave the way for more big chains" in the city which would "destroy independently-run businesses and threaten its cultural identity".
James Anderson, associate director at planning consultancy Turley, which is leading community engagement on the plans, said Glasfryn Road was an "important development for St Davids" and "one the joint applicants want to get right".
"The issues raised by the petition will be clarified at the public exhibitions and the full team will be available to explain the proposals and how the new homes and hotel are working together," he said.
"Feedback on the latest proposals is important to us and we are looking forward to talking residents through the latest plans and listening to their views."
The exhibitions will be held at Oriel y Parc in St Davids on Friday and Saturday.


This week Pyongyang is a city in a frenzy of preparation.

At times the crowds of people thronging the sidewalks have turned the streets into a blaze of colour - the women, vivid explosions of rainbow hues in their traditional Korean dresses, and the men, although black or grey suited, carrying large artificial pink and red flowers.
These plastic, pom-pom like azaleas are used for waving in the ritualised adulation of their leader for which they are now so busy rehearsing.
Some sit in large groups waiting for instructions, others walk purposefully to or from the parade ground, chatting or laughing together along the streets of a capital city that is still largely devoid of traffic.
Antiquated army trucks with open tops trundle into town, in convoys dozens long, each packed with soldiers in uniform.
There's a relaxed, holiday feel: the female conscripts, separated in their own trucks, smiling and waving at passers-by, the male troops singing merrily in unison.
It gives this city the air of a giant film set for a war-time period costume drama.
But in the world's last truly totalitarian state, this is the reality.
Mass mobilisation is the defining essence of social and political life and there is no more important occasion for the expression of absolute fealty to the leader than The Day of the Sun.
Saturday marks the 105th anniversary of the birth of North Korea's long-dead founding president Kim Il-sung, although according to the country's constitution he remains formally in office.
The celebrations this year have taken on an added sense of symbolic meaning as they take place amid one of the periodic peaks in the tension that has so often defined the country's relationship with the outside world.
This isolated regime has shown a skilful knack of posting itself into the priority inbox of every US president who has come to office in recent years.
This time is no different, with an underground nuclear test shortly before the election, followed by a flurry of ballistic missile tests during the first few weeks of Mr Trump's presidency.
And like other presidents before him, Mr Trump appears to be exploring whether - instead of the failed diplomacy, debate and delay - a more direct, dramatic option might be available.
Whether he will, like others, finish his term having eventually decided that there is no realistic alternative remains to be seen.
But the fact that no US administration has yet been able to find a way to rein in North Korea's nuclear ambitions is a measure of the remarkable success of its game of brinkmanship with the outside world.
It is a classic study in military deterrence.
The regime's most powerful weapon has long been its conventional artillery.
Placed close to the border, it could cause significant damage and large loss of life in the South Korean capital within a matter of minutes.
For the government in Seoul it makes not just any offensive military option unthinkable, but complicates even defensive calculation.
In 2010, following an audacious and unprovoked torpedo attack on the South Korean warship the Cheonan - claiming the lives of 46 sailors and widely believed by the international community to have been carried out by the North - the South sat on its hands.
A few months later there was also no retaliation when North Korea shelled a South Korean island, hitting both military and civilian targets.
It is proof that the North has calculated all too well the costs of military engagement for its democratic, populous and economically vibrant neighbour.
Employing exactly the same logic, Pyongyang has been edging ever closer to possessing a deliverable nuclear arsenal with the aim of forcing its foes further afield into the same strategic bind.
The one thing that keeps the North Korean leadership awake at night is the thought of the B52 bombers stationed on the Pacific island of Guam.
And they have learned a very particular lesson from the US-led efforts to bring about regime change elsewhere in the world.
Iraq didn't have nuclear weapons and Libya had given its up.
Unless North Korea can be given the kind of guarantees that would make it feel secure enough - which seems unlikely in the short term - then any effort to negotiate away its nuclear programme is bound to fail.
So for now, the world is left with two stark choices.
Accept North Korea as a member of the nuclear-armed club, or try to force it to disarm, either through ever tougher sanctions or the incalculably risky option of military action.
As President Trump weighs these options, North Koreans are preparing to march, dance and sing to the glory of the country's ruling family this weekend.
And there is speculation that another nuclear test could be a matter of just days away.


The ex-husband and father of an alleged "honour killing" victim have been remanded in custody for a further 15 days by a judge in Pakistan.

Samia Shahid, 28, from Bradford, died in the country in July.
Chaudhry Muhammad Shakeel is accused of her murder while her father Chaudhry Muhammad Shahid is being held as a suspected accessory to the crime.
The pair will next appear in court in Jhelum, in the northern Punjab province, on 26 November.
Both were returned to prison following a request by a lawyer acting for Ms Shahid's second husband, Syed Mukhtar Kazim, for more time to review the case.
Neither Mr Shakeel nor Mr Shahid have been formally charged and their lawyers have previously argued there is no evidence against them.
Last month Ms Shahid's mother, Imtiaz BiBi, and sister, Madiha Shahid, were both declared proclaimed offenders in Pakistan meaning police believe they were involved in her death and want to question them.
Arrest warrants were issued by a judge when they did not appear in court.
Police are applying for a red notice - an international alert for a wanted person - from Interpol. The process is expected to take weeks or even months before one is issued.
The BBC understands extradition requests have not yet been made to the UK authorities.
Ms Shahid, a beautician, married Mr Kazim in Leeds in 2014 and the couple moved to Dubai.
Mr Kazim has claimed his wife, who died while visiting relatives in Pakistan, was killed because her family disapproved of their marriage.
Initially it was claimed she had died of a heart attack but a post-mortem examination found she had been strangled.


An overhauled state pension - being paid to new, rather than existing pensioners - has begun, with some set to gain while others lose out.

The government's long-term aim is simplification, stripping away extras such as the second state pension.
It will also become cheaper, in time, for the government with many of those in their 20s and 30s receiving less than they would under the old system.
But in the short-term, the self-employed particularly will benefit.
A Department for Work and Pensions (DWP) spokeswoman said: "Millions stand to gain from the changes to the state pension, including women and the self-employed who so often lost out in the past.
"The new state pension will provide a sustainable system for future generations who will also benefit from workplace pension savings throughout their careers."
However, the independent Institute for Fiscal Studies (IFS) said that the new single tier, or flat-rate, element of the new system may have been overstated, leading to the risk of "disillusionment" from new pensioners.
"The aim is to replace the complex mess of existing rules with a new, far simpler system that rewards a wide range of contributions - whether that be paid employment or caring for children - in exactly the same way," said Rowena Crawford, of the IFS.
"We estimate that women will gain on average £5.20 per week in additional state pension income at the state pension age, and those who have been self-employed for at least 10 years will gain an average of £7.50 per week.
"But continued complexity is unavoidable in the short-run. There is a considerable risk of disillusionment as people start claiming pension incomes this year."
She said that some would receive a "nasty surprise" when they realised they would receive less than the advertised "flat-rate" of £155.65 a week.
For more information about whether you are a winner or a loser under the new system, click here.
The basic state pension, which is still being paid to existing pensioners, is worth about £120 a week, plus top-ups such as the state second pension for those who qualify.
The new state pension will be paid to men born after 6 April 1951 and women born after 6 April 1953.
To qualify for any state pension under the new system, individuals will need to have 10 years of National Insurance contributions. To receive the full amount, of £155.65, they will need to have 35 qualifying years.
Some, primarily public sector workers, will receive less than the full amount if they were contracted out of the state second pension. In practical terms, this meant they paid less in National Insurance contributions, but more into their workplace pension scheme.
The most significant change in the long-term is the abolition of the state second pension.
The effect, as revealed by BBC News earlier in the week, is that 11.4 million younger workers will get less out of the new system than they would have done had the old system carried on, according to the Pensions Policy Institute.
By 2050, about half of retirees will get a higher pay-out, with half getting a lower pay-out.
Assuming a pension age of 70 by then, in general, people born before 1980 can, on average, expect to do better out of the new system, but those born after that date are likely to fare worse.


Laura Trevelyan describes what it was like to report on Hurricane Sandy.

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"Reporting a hurricane is an alarming experience, it starts when you're on the highway on the way to the story and you see the big signs overhead that say "State of Emergency Evacuation Ordered" and you're actually going against the traffic.
Everyone's leaving town and you're heading into the heart of a hurricane, it doesn't feel like the most sensible thing you should be doing, but that's where the story is so that's where you go.
And then when you get to the water's edge, there are roadblocks everywhere, the police are telling you that everyone has gone, but you have a press pass, so you can cross the police lines. Then you see the water, the waves are 20 feet high, they were when I was in New Jersey, when Atlantic City was evacuated, and again everybody is going in the opposite direction to you.
People are rain sodden, they are carrying all their possessions, their animals, they're all leaving town and you're going further into the heart of the hurricane.   And just before the hurricane makes land fall and you're outside and the trees are blowing backwards, your hair is blowing in your face, the water is horizontal, driving into your eyes you can't even see.
Then, you see that all of the birds have gone, everything is gone you seem to be the last thing left.   You take refuge in the news truck and it begins to rock as the winds reach 60 or even 80 miles per hour and you wonder, is this safe, is this sensible, is this what my family think that I should be doing.  But at the same time it's of such importance to report on the progress of this storm that you stay put, and you do your job and you live the story. "
Read Laura's report on Hurricane Sandy
In pictures:  New York flooded by Hurricane Sandy
Read the BBC's coverage of Hurricane Sandy
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With journalists in more countries than any other international news broadcaster, we don't just report a story, we live it.
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Katrina Esau is working hard to save the language of her childhood from dying out.

At 84, Ms Esau is one of the last three fluent speakers of N||uu, one of the languages spoken South Africa's San community, also known as Bushmen.
N||uu is considered the original language of southern Africa.
With no other fluent speakers in the world apart from this family, the language is recognised by the UN as "critically endangered".
"When I was a child, I only spoke N||uu and I heard a lot of people speaking the language. Those were good times, we loved our language but that has changed," says Ms Esau in Upington, a town in the Northern Cape Province.
For centuries, the San roamed this region freely, gathering plants and hunting animals to feed their families.
But today the traditional practices of the San have all but died out and their descendants tell me that language is one of the only things left that connects them to their history.
Inside a small wooden hut, she teaches the 112 sounds and 45 distinct clicks of N||uu with the local children.
"I'm teaching the language because I don't want it to become extinct when we die," Ms Esau says.
"I want to pass as much of it as I can but I am very aware that we don't have a lot of time."
Ms Esau has been running the school in her home for about 10 years.
The people in this community, including Ms Esau now mainly speak Afrikaans - a language brought by the Dutch settlers who arrived in South Africa in the 17th Century.
"We would get beaten up by the white man if we were caught speaking our language," she tells me.
"Because of our history, people today do not want to speak the language any more, there is so much pain around it.
"We abandoned the N||uu language and learned to speak Afrikaans, although we are not white people - that has affected our identity," she adds.
Ms Esau's two sisters Hanna Koper and Griet Seekoei - both over 95 - are listening intently as she speaks with bitter fondness of their childhood.
They don't speak much but nod in agreement as she speaks.
Ouma Geelmeid, as she is affectionately known, says she is hoping to remove the shame around speaking N||uu today.
During lessons, with a stick in her hand, Ms Esau points out the N||uu names for body parts on the white board as the students read in chorus.
Like many other African languages, this language had been passed down orally over generations - but this is now threatening its survival.
Until recently years there was no record of it as a written language.
Ms Esau worked with linguists, Sheena Shah from the School of Oriental and African Studies (Soas) in London and Matthias Brezinger of the The Centre for African Language Diversity Centre in Cape Town to create an N||uu alphabet and basic rules of grammar.
"From the work we did with Ouma Geelmeid's community, we learned that these communities see language as their identity and that is an important aspect to human interactions." says Ms Shah.
"Personal identity is incredibly important, more so now in a global world."
Language is about more than simply an idea to communicate with one another, it is also tied in with culture and a way of life for a community, the experts tell me.
"When you look at the African languages, you learn that they help communicate different perspectives on life, relationships, spirituality, the earth, health, humanity," says Mr Brezinger.
"There is a wealth of knowledge on survival that has been passed down through the years in indigenous communities that the Western world knows very little about and when these languages die, that unique knowledge is also lost," he continues.
Back in the classroom, there are about 20 children, most of them under 10 years old and a few teenagers.
Mary-Ann Prins, 16, is Ms Esua's best student and hopes to one day teach this class.
"I love learning this language. It makes me feel like I belong, like I am connected to my great-grandparents. I'm told that they used to speak it and today I can be a part of that to," she says with a broad smile.
The N||uu is not the only language at risk of disappearing in South Africa.
Three hours away, in the town of Springbok, Nama speakers have been lobbying the government to have their language made an official language.
Despite being historically widely spoken in South Africa, Nama is not recognised as one of the country's 11 official languages.
"It's very sad that our children cannot speak Nama. It breaks my heart that our children will never be able communicate with their elders in their own language," says Maria Damara, 95, one of the only Nama speakers here.
"What future will they have, what will happen to our culture?"
South Africa's top six mother-tongue languages:
Source: SA.info/Census 2011
Why SA students want to be taught in English
Community leader Wiela Beker, 56, agrees:
"If you don't have language, you don't have nothing. I'm talking in English to you now but I am not English. I want to speak Nama because that's what and who I am.
"Unless we do something about it, our culture is going to die. We are fighting for our culture when we fight for our language," he says.
But without a shift in language policy and government intervention, this community say they are worried that it won't be long until their language finds itself in the same boat at as N||uu - on the verge of extinction.
And so for these communities the fight is for a shared identity, a sense of belonging - and therefore a legacy for future generations


Donald Trump's campaign for the US presidency has been thrown into turmoil by a video carrying obscene remarks about women.

In the video, Mr Trump says "you can do anything" to women "when you're a star" and brags about trying to grope and kiss women. He has since apologised.
Mr Trump has clearly been an unpredictable candidate but no previous revelation or off-the-cuff comment has generated this much reaction.
So who in the Republican camp is still backing him and who has decided enough is enough?
The Utah governor, who previously supported Mr Trump, tweeted that the comments he had made were "beyond offensive and despicable". He continued: "While I cannot vote for Hillary Clinton, I will not vote for Trump."
A congressman for Utah, he withdrew support live on television.
He said: "It's sad really, but I can't endorse Donald Trump for president after those comments and the way he said them.
"I can't look my 15-year-old daughter in the eye and tell her I endorse this man to become president."
Like his Utah colleague Mr Herbert, he said the video left him at a loss regarding who to vote for, as there was "no way" he would vote for Hillary Clinton.
Mr Chaffetz is the chairman of the committee which is investigating Mrs Clinton's use of emails while she was secretary of state.
She was the only woman who ran for the Republican candidacy for 2016 and was briefly the vice-presidential candidate on Ted Cruz's ticket.
Although she once said she was "horrified" by Mr Trump, in September she finally gave him her support - but only as an alternative to Mrs Clinton.
In response to the tape, she said: "We must have a conservative in the White House to restore accountability, opportunity and security. For the sake of our Constitution and the rule of law, we must defeat Hillary Clinton. Today I ask Donald Trump to step aside and for the RNC to replace him with Gov Mike Pence."
The prominent senator for Arizona and one-time presidential nominee said Mr Trump should "suffer the consequences" of his remarks.
"Donald Trump's behaviour make(s) it impossible to continue to offer even conditional support for his candidacy," he said.
"There are no excuses for Donald Trump's offensive and demeaning comments.
"No woman should ever be victimised by this kind of inappropriate behaviour. He alone bears the burden of his conduct and alone should suffer the consequences."
The former secretary of state wrote on her Facebook page: "Enough! Donald Trump should not be President. He should withdraw.
"As a Republican, I hope to support someone who has the dignity and stature to run for the highest office in the greatest democracy on Earth."
The former California governor, who is more famous for his acting career and who took over from Mr Trump as a host of The Apprentice on NBC, said he would not vote for his predecessor in the presidential election.
He said: "I want to take a moment to remind my fellow Republicans that it is not only acceptable to choose your country over your party - it is your duty."
The former governor of New York tweeted that he was "horrified" at the tape and that Mr Trump's campaign was "a poisonous mix of bigotry and ignorance".
"Enough!" Mr Pataki tweeted. "He needs to step down."
A former governor of Minnesota, he said the comments were "unacceptable, and disqualifying for someone who hopes to serve as Commander in Chief".
A former Utah governor, Mr Huntsman called for the Republican vice-presidential candidate to take over running for president instead.
"In a campaign cycle that has been nothing but a race to the bottom, at such a critical moment for our nation, the time has come for Governor Pence to lead the ticket," he told the Salt Lake City Tribune in his home state.
Another Utah Republican, the senator addressed Mr Trump with the words: "With all due respect sir, you, sir, are the distraction. Your conduct, sir, is the distraction. It's a distraction from the very principles that will help us win in November."
He continued: "I respectfully ask you, with all due respect, to step aside, step down, allow someone else to carry the banner of these principles, these principles that have made our country great, these principles that will stand as a beacon of hope to the American family."
The Colorado congressman was one of the first Republicans to call on Mr Trump to stop running for president after the tapes of his comments were broadcast.
On Friday, he said: "For the good of the country, and to give the Republicans a chance of defeating Hillary Clinton, Mr. Trump should step aside.
"His defeat at this point seems almost certain."
An Alabama senator, she said: "As disappointed as I've been with his antics throughout this campaign, I thought supporting the nominee was the best thing for our country and our party.
"Now, it is abundantly clear that the best thing for our country and our party is for Trump to step aside and allow a responsible, respectable Republican to lead the ticket."
A New Jersey congressman, he said: "I cannot support and will not vote for Donald Trump or Hillary Clinton to be president of the United States. I will write in Governor Mike Pence for President."
The Virginia congresswoman, who had not yet declared support for Mr Trump, said she could not "in good conscience" vote for him.
She echoed calls for somebody else to replace him, saying: "This is disgusting, vile, and disqualifying.
"No woman should ever be subjected to this type of obscene behaviour and it is unbecoming of anybody seeking high office.
"In light of these comments, Donald Trump should step aside and allow our party to replace him with Mike Pence or another appropriate nominee from the Republican Party."
A senator for Alaska who had not yet declared her support for Mr Trump either, she said she watched the video immediately after a meeting about sexual exploitation and she felt "disgusted".
She said Mr Trump had "forfeited the right" to be the Republican party's nominee.
A Nebraska senator, she said: "The comments made by Mr Trump were disgusting and totally unacceptable under any circumstance. It would be wise for him to step aside and allow Mike Pence to serve as our party's nominee."
The Idaho senator also called for another candidate to be put forward instead, and said: "I have reached the decision that I can no longer endorse Donald Trump. This is not a decision that I have reached lightly, but his pattern of behaviour has left me no choice."
A Utah congressman, he said: "I'm incredibly disappointed in our party's candidate. And unlike the Democrats who have proven completely unwilling to hold secretary Clinton accountable for her illegal activities that endangered our national security, I am willing to hold Mr Trump accountable.
"I am therefore calling for him to step aside and to allow Mike Pence to lead our party."
An Arizona senator, he tweeted that "America deserves far better" than Mr Trump and that he should withdraw from the race.
A Nebraska congressman, he called for Mr Trump to step aside "and allow Mike Pence to become the Republican nominee".
A New Jersey congressman, he said: "Donald Trump's comments are inexcusable. I am appalled that he would brag about violating a woman's physical boundaries.
"As a husband and father of two daughters, I denounce his comments and the behaviour that it incites. I believe that Mike Pence would be the best nominee for the Republican Party to defeat Hillary Clinton."
A Nevada congressman, he had previously said he was "100%" behind Mr Trump.
But on Saturday, he pulled his support, saying: "I think that when we degrade that mother, wife, housewife, whatever you want to deal with, daughter - that you degrade America."
Another Nevada congressman, he said he was "disappointed in our choices for president" and did not support Mr Trump any more.
He said: "I can no longer look past the pattern of behaviour and comments that have been made by Donald Trump."
The governor of Alabama, who had previously endorsed John Kasich, said: "I cannot and will not vote for Donald Trump."
The governor of Nevada said: "This video exposed not just words, but now an established pattern which I find to be repulsive and unacceptable for a candidate for President of the United States."
The governor of South Dakota said: "Enough is enough. Donald Trump should withdraw in favour of Governor Mike Pence. This election is too important."
The Minnesota congressman said the comments were "disgusting and offensive" and he would not vote for Mr Trump.
A senator in Alaska, he said: "Keeping Republicans in the Senate majority is critical to the economic and national security of Alaska and America.
"As for the White House, Donald Trump should step aside. I will support Mike Pence for president."
An Alabama congressman, he said: "There are absolutely no circumstances under which it would ever be appropriate to speak of women in such a way.
"It is now clear Donald Trump is not fit to be president of the United States and cannot defeat Hillary Clinton."
The Michigan congressman said Mr Trump should "carefully consider stepping aside from the ticket".
He said: "I urge him to think about our country over his own candidacy."
An Illinois congressman, he took the rare step of referencing his sons when explaining why he would not support Mr Trump any more. Many others had referenced their female relatives.
In a statement, he said: "As parents of a teenage daughter and teen twin boys, my wife and I teach them to respect women and that they will be judged by their words and actions.
"The abhorrent comments made by Donald Trump are inexcusable and go directly against what I've been doing in Washington to combat assaults on college campuses. Because of this, I am rescinding my support for Donald Trump and asking to have my name removed from his agriculture advisory committee."
The Florida congressman originally supported Marco Rubio for the presidential nomination, but then called for party unity behind Mr Trump.
Following the publication of the tape, he said: "My greatest responsibility in life is to try and be a good husband and father. If I support him for president, I will be telling my boys that I think it's okay to treat women like objects - and I'll have failed as a dad. Therefore, I can no longer support Donald Trump for president and will not be voting for him or Hillary Clinton."
The Missouri congresswoman said: "As a strong and vocal advocate for victims of sex trafficking and assault, I must be true to those survivors and myself and condemn the predatory and reprehensible comments of Donald Trump."
The New Hampshire senator, who is running for re-election in November, withdrew her support from Mr Trump and said she would write Mike Pence's name on her ballot paper instead.
She said: "I'm a mom and an American first, and I cannot and will not support a candidate for president who brags about degrading and assaulting women.
Also facing a re-election battle, this Ohio senator said: "While I continue to respect those who still support Donald Trump, I can no longer support him."
Mr Glenn is running for the Senate. From Colorado, he asked Mr Trump to "do the honourable, selfless thing" and step aside to allow Mr Pence to run for President.
In a statement, he wrote: "If Trump is truly committed to making America great again, then this is the only way forward.
"As a father, as a Christian, and as a Republican, I believe that we simply cannot tolerate a nominee who speaks this way about women."
The chairman of the Republican Conference and senator for South Dakota said: "Donald Trump should withdraw and Mike Pence should be our nominee effective immediately."
The former secretary of education served under Ronald Reagan. He is now a conservative commentator and it was only in August that he said Republicans who opposed Mr Trump "suffer from a terrible case of moral superiority and put their own vanity and taste above the interest of the country".
But after seeing the video, he too called on Mr Trump to stand down, saying: "It's over. I hate to say it, but it's over."
"It's a shame, a crying shame, but he can't win," he said.
The House speaker, who endorsed Mr Trump just a few weeks ago, has withdrawn an invitation for the presidential candidate to attend the Republican Fall Fest in his home state of Wisconsin this weekend.
Mr Ryan has not pulled his endorsement, but it does pack a powerful punch for a candidate to be uninvited from an event just one month before the election.
Mr Ryan said: "I am sickened by what I heard today. Women are to be championed and revered, not objectified. I hope Mr Trump treats this situation with the seriousness it deserves and works to demonstrate to the country that he has greater respect for women than this clip suggests. In the meantime, he is no longer attending tomorrow's event in Wisconsin."
The Senate majority leader said the comments were "repugnant", adding that Mr Trump "needs to apologise directly to women and girls everywhere".
He also said that Mr Trump should "take full responsibility for the utter lack of respect for women shown in his comments on that tape".
The Utah state house speaker said he hoped for an apology from Mr Trump but did not rescind his support.
He said: "To say I'm disappointed would be a gross understatement."
The former Florida governor, who ran unsuccessfully for president this year, tweeted: "As the grandfather of two precious girls, I find that no apology can excuse away Donald Trump's reprehensible comments degrading women."
He did not mention his cousin Billy Bush, who was Mr Trump's interlocutor in the incriminating video.
The former presidential candidate has been critical of Mr Trump in the past and said he would not vote for him.
After the video emerged, he tweeted that the comments the current candidate made were "vile degradations" with an impact on women and on the image of the US around the world.
He wrote: "Hitting on married women? Condoning assault? Such vile degradations demean our wives and daughters and corrupt America's face to the world."
The California congressman said: "In my career as a law enforcement professional I was confronted with and worked tirelessly to end the horrific reality of violence toward women.
"After hearing Donald Trump's inexcusable comments I was deeply disturbed & reminded of that reprehensible behaviour.
"While I've never before endorsed a Presidential candidate, I've felt compelled to strongly condemn many of Mr. Trump's previous outrageous remarks. And after serious consideration, I have decided that I cannot support either candidate for President."
The governor of New Mexico had never endorsed Mr Trump. She said: "What Trump brags about is appalling and completely unacceptable.
"No woman should ever be treated the way he claims he treated women. Unfortunately, there is a pattern of disturbing conduct and offensive rhetoric that raises serious questions about his fitness to be President.
"That's why I have withheld my support from the very beginning, and will not support him now."
The Ohio governor, who also ran for the presidency and was beaten by Mr Trump, had already said he would not vote for him. He tweeted his condemnation of Mr Trump's comments, with the words: "Make no mistake the comments were wrong and offensive. They are indefensible."
A congresswoman from Utah, where much of this wave of condemnation of Mr Trump began, she said: "For the past several months I have been one of the few who refused to endorse Donald Trump.
"I have said all along that I was still waiting for Mr Trump to demonstrate his commitment to the kinds of principles and policies the people in Utah's 4th Congressional District want in their elected leaders. Mr Trump has yet to clear that bar and his behaviour and bravado have reached a new low.
"I cannot vote for him. For the good of the party, and the country, he should step aside."
The senator for Colorado called on Mr Trump to stand down, calling him "a candidate whose flaws are beyond mere moral shortcomings and who shows a disgust for American values and a disdain for dignity unbecoming of the Presidency".
The lieutenant-governor of New Jersey had not endorsed Mr Trump, despite working closely in her home state with his prominent supporter Chris Christie.
In response to a question on Twitter, she wrote: "No apology can excuse away Mr. Trump's reprehensible comments degrading women. We're raising my three boys to be better than that."
She later said she would vote, but not for Mrs Clinton and not for Mr Trump.
The Pennsylvania congressman was another Republican politician who had declined to endorse Mr Trump, deflecting a question when asked his stance back in May. After the 2005 video surfaced, he called for Mr Trump to "end his candidacy" and said: "This sort of vile talk is appalling, it's offensive, and there's no place in public or private for it. It's simply wrong.
The Governor of Tennessee called on Mr Trump to stand aside and make way for Mr Pence. He said he would write another Republican's name on the ballot paper otherwise, and added: "Character in our leaders does matter. None of us in elected office are perfect, but the decisions that are made in the Oval Office have too many consequences to ignore the behaviour we have seen."
In a text message to his local newspaper, the Texas congressman wrote: "I never endorsed Trump and I cannot in good conscience support or vote for a man who degrades women, insults minorities and has no clear path to keep our country safe. He should step aside for a true conservative to beat Hillary Clinton."
A congressman from New York, he has previously disavowed Mr Trump's "tone and rhetoric" but after the video, told his local newspaper that "what he was talking about is tantamount to sexual assault".
Mr Katko called on Mr Trump to stand aside and let somebody else run, saying:  "I've talked repeatedly with my family and people I consider close to me. We are all roundly disgusted.
"We just all came to the same conclusion that he does not deserve support. You just can't explain this away."
A congressman for Idaho, he said: "While I've never endorsed Donald Trump, I find his recent comments about women deplorable. In my opinion, he has demonstrated that he is unfit to be president and I cannot support him."
The only woman who represents Texas in the US Congress had previously declined to endorse Mr Trump. After the video surfaced, she said: "We have heard rumours about the insensitive and vulgar things Mr. Trump says about women. But watching that video is disgusting. Mr. Trump should remove himself from consideration as Commander in Chief."
A Washington congresswoman, she said she will write in Paul Ryan's name on her ballot paper, instead of voting for Mr Trump. In an email, she wrote: ""For months I've left the door open for Donald Trump to earn my vote. That door has now slammed shut."


Banbridge will contest the European Hockey League knockout round of 16 for the first time after a dramatic 3-3 draw with Belgian side Royal Leopold.

Both sides had defeated Saint Germain from France in their opening pool game but the 2-0 win for Bann on Saturday edged them ahead on goal difference.
The Ulster side, who hosted the Round One tournament at Havelock Park, will join Lisnagarvey in the knockout stage.
That tournament will be held next April in the Netherlands.
Bann got off to a dream start on Sunday, Jonny McKee picking out Eugene Magee and the Rio Olympian slotted home on the reverse stick to open the scoring within ninety seconds of the game getting underway.
Royal Leopold, who included two-time World Cup winner Glenn Turner from Australia in their line-up, regrouped quickly and by half-time had taken the lead converting two penalty corners from Max Plennevaux and Alexis Lemaire.
But for the brilliance of Bann goalkeeper Gareth Lennox it could have been worse.
However the home side, backed by a crowd of over two thousand, showed their character early in the second half as Kyle Marshall slid on to a Matthew Bell push from a corner to deflect the ball into the net to tie the score at 2-2.
Almost immediately Dorian Thiery stole in to lob Lennox to restore the advantage for the Belgians but Marshall once again got on the end of a Bell pass to make it 3-3 with 21 minutes to go.
The final quarter was a tense affair as Leopold pushed for the winning goal but with Bell marshalling the defence superbly and Lennox producing a string of good saves, Banbridge held on to record a famous result.
The other three places in the knockout 16 went to Spaniards Atletic Terrassa, UHC Hamburg and defending champions Kampong from Holland, whose goalkeeper is Ireland international Davy Harte.
The Dutch outfit qualified at the expense of local GB international Iain Lewers and his club side Holcombe.


Captain Wayne Rooney has "unreservedly" apologised to interim England manager Gareth Southgate after pictures emerged showing him at a wedding at the team's hotel while on international duty.

A statement issued on behalf of Rooney, 31, called the images "inappropriate".
The Sun published images it said showed Rooney drunk at a party late on Saturday night - a day after England's World Cup qualifying win over Scotland.
The FA, meanwhile, says it will review its policies around players' free time.
Thursday's edition of The Sun claims that policy review was launched because 10 England players were at a nightclub until 04:30 GMT on Sunday.
The statement from Rooney's management continued: "Naturally Wayne is sorry that pictures taken with fans have been published.
"Although it was a day off for the whole squad and staff, he fully recognises that the images are inappropriate for someone in his position.
"Earlier today Wayne spoke privately to both Gareth Southgate and [FA technical director] Dan Ashworth to unreservedly apologise.
"He would like to further extend that apology to any young fans who have seen these pictures."
Rooney missed Tuesday's 2-2 friendly draw with Spain with a minor knee injury.
Rooney's apology came as the FA issued a statement saying "England personnel have a responsibility to behave appropriately at all times".
The governing body added: "We will be reviewing our policy around free time while on international duty."


A former priest has been found guilty of indecently assaulting a secondary school student in the 1980s in the Republic of Ireland.

Henry Moloney worked at the time as a choirmaster and music teacher at Rockwell College, near Cashel, County Tipperary.
The 77-year-old, of Kimmage Manor, Dublin, was found guilty of seven counts of indecent assault on the schoolboy.
He had denied the charges.
The victim had told Clonmel Circuit Court how the abuse began near the start of a school year after Moloney asked him to join the choir.
He said although he knew what had happened was wrong, he thought no-one would believe him instead of a priest.
The offences all took place in the course of one school year, in the college's chapel and in Moloney's room there.
Moloney is due to be sentenced on 15 December.


Super-middleweight George Groves will take on Fedor Chudinov for the WBA world title at Bramall Lane on 27 May.

Groves, 29, has lost in contesting world honours three times, twice to Carl Froch and most recently to Badou Jack, for the WBC strap he has since vacated.
Chudinov, also 29, held the WBA title until defeat by Felix Sturm in February of last year.
But Sturm vacated the title in October having tested positive for a steroid.
The controversial points defeat was the first of Chudinov's 15-fight career, while Groves will enter the Sheffield bout with a record of 25 wins and three losses.
In December, Groves retained his WBA International super-middleweight title with a unanimous points win over Eduard Gutknecht.
The contest will form part of the undercard as Sheffield's Kell Brook defends his IBF world welterweight title against undefeated American Errol Spence Jr.


It is fair to say that entrepreneur Simon Lee has a flair for languages.

The 32-year-old South Korean speaks no less than six of them - Korean, English, French, Arabic, Mandarin and Japanese.
And he says he plans to learn more.
With such linguistic skills Mr Lee doesn't have much personal need for translation services, but after friends at university in Seoul started asking him for help with their English homework he was inspired to set up just such a company.
Launched in September 2012, his business - Flitto - now has more than five million users around the world, and enjoys revenues of about $2.1m (£1.4m) a year.
Unlike most professional translation companies, it doesn't directly employ translators.
Instead, Flitto invites members of the public who know more than one language to offer their services via its website and mobile phone app.
Today it has one million translators in 170 countries on its books, who offer translations between 17 different languages. They all get paid for each piece of work they do, from which Flitto takes a small percentage.
In the first feature of new six-part series called "The Making of Me", we look back at key stages in Mr Lee's life and the development of his business.
Due to his father's job with a global company, Mr Lee spent most of his childhood living overseas.
He was born in Kuwait, where he spent his first four years, before the family then moved to UK.
After three years in the UK, the family relocated again, this time to the US, before moving to Jeddah in Saudi Arabia 24 months later.
Educated in international schools, he was quick to pick up English, French and Arabic, the first three languages Mr Lee added to his native Korean.
"I met all these different people from different countries," he says.
"What I realised is that all human beings are the same you know, but because of the different languages we misunderstand each other."
After seven years in Saudi Arabia, Mr Lee's family moved back to South Korea where he finished high school, and then went to the country's prestigious Korean University.
It was at college that friends of his started to ask him to help them translate their homework into English, which was required as part of their courses.
"So friends would say to me, 'Simon, if you translate this homework into English I'm going to buy you dinner'," he says.
"And more and more friends were asking me to do that, so I didn't have enough time."
Realising the business potential of translation services, Mr Lee started getting other multilingual friends to help out, and his idea for setting up a company was born.
Flitto was launched in September 2012, but rather than start the business in South Korea, Mr Lee decided to relocate to London.
He said he chose to do this for two main reasons - he wanted to temporarily get away from his friends who had well paid jobs in corporate South Korea, and because London is such a multi-cultural city.
He says: "I was starting a business, so I had no money. So what I thought is that [if I stayed in Seoul] I would compare myself with my friends, and that would make me really sad.
"So I just wanted to go some place quiet. And London is the centre of Europe, with lots and lots of different people who speak different languages."
To help get the business up and running, Mr Lee based himself at a shared working space in London run by a start-up support organisation called Techstars. This gave him both mentoring support and funding.
Yet with little funds to advertise, Mr Lee had to come up with a novel way of drawing attention to Flitto's website and app, and attracting both translators and customers.
His answer was for Flitto to collate the Twitter and other social media feeds of Western pop stars such as Lady Gaga, and encourage people to translate them into different languages, in exchange for merchandise of the celebrity in question.
It worked, and Flitto soon started to see its business and brand recognition grow, with more and more customers, and a growing number of translators signing up.
Today Flitto, which has its headquarters in Seoul, and has just 34 full-time staff, says it gets 70,000 translation requests per day.
These range from individuals wanting a few lines translated, to businesses requesting bulk work. Each translator - all one million of them - is graded out of five in terms of feedback on the quality of their work.
As an additional revenue stream Flitto's website also has a store section, where people can buy everything from clothing to iPhone cases. And to further drive traffic to the site, it also has an editorial section, which includes stories and photo galleries from around the world.
Mr Lee says the fact that Flitto doesn't directly employ translators means that it can significantly undercut the traditional translation service providers. Its costs start from 10 cents (6p) for 250 characters.
And he says that he does not feel threatened by the growth of online computerized translation services because he does not believe they will ever be as accurate as human beings doing the work.
Mr Lee says his biggest error was an earlier attempt to set up a similar translation business.
This was back in 2007, when he launched a company called Flyingcane.
He said it was the right idea, just at the wrong time.
"The problem was that there were no smart phones back in 2007," he says.
He said that as a result of this, there was obviously no mobile phone app side to the business, and this meant that the translation work was slower because people couldn't do it as easily when they were on the move.
After Flyingcane failed, Mr Lee then spent a number of years working for a South Korean mobile phone network, before returning to his business idea in 2011, and then launching Flitto in 2012.
In terms of the day-to-day running of the company, Mr Lee says he can be a workaholic, and didn't take a holiday for six years.
In the end, matters came to a head with fellow board members.
He says: "My investors forced me to take a vacation, which I never wanted. They told me 'Simon, you don't look good, you don't look healthy, so we think you need to time to go off for fresh air, and to relax yourself'."
Mr Lee says that people thinking of setting up their own businesses need to be aware that it can take it out of you.
"Whenever my juniors ask me 'Simon do you recommend that I should start my own business?', I say 'no, there is too much stress'."
But despite the inevitable anxiety that running your own company brings, Mr Lee says he recommends that people try it.
"Just stick to your dream and believe in yourself, that's my advice.
"Everyday you feel your heartbeat, that you are actually living your life."


If one human evolution paper published in 2013 sticks in my mind above all others, it has to be the wonderful report in the 18 October issue of the journal Science.

The article in question described the beautiful fifth skull from Dmanisi in Georgia. Most commentators and colleagues were full of praise, but controversy soon reared its ugly head.
What was, in my view, a logical conclusion reached by the authors was too much for some researchers to take.
The conclusion of the Dmanisi study was that the variation in skull shape and morphology observed in this small sample, derived from a single population of Homo erectus, matched the entire variation observed among African fossils ascribed to three species - H. erectus, H. habilis and H. rudolfensis.
The five highly variable Dmanisi fossils belonged to a single population of H. erectus, so how could we argue any longer that similar variation among spatially and temporally widely distributed fossils in Africa reflected differences between species? They all had to be the same species.
I have been advocating that the morphological differences observed within fossils typically ascribed to Homo sapiens (the so-called modern humans) and the Neanderthals fall within the variation observable in a single species.
It was not surprising to find that Neanderthals and modern humans interbred, a clear expectation of the biological species concept.
But most people were surprised with that particular discovery, as indeed they were with the fifth skull and many other recent discoveries, for example the "Hobbit" from the Indonesian island of Flores.
It seems that almost every other discovery in palaeoanthropology is reported as a surprise. I wonder when the penny will drop: when we have five pieces of a 5,000-piece jigsaw puzzle, every new bit that we add is likely to change the picture.
Did we really think that having just a minuscule residue of our long and diverse past was enough for us to tell humanity's story?
If the fossils of 1.8 or so million years ago and those of the more recent Neanderthal-modern human era were all part of a single, morphologically diverse, species with a wide geographical range, what is there to suggest that it would have been any different in the intervening periods?
Probably not so different if we take the latest finds from the Altai Mountains in Siberia into account.  Denisova Cave has produced yet another surprise, revealing that, not only was there gene flow between Neanderthals, Denisovans and modern humans, but that a fourth player was also involved in the gene-exchange game.
The identity of the fourth player remains unknown but it was an ancient lineage that had been separate for probably over a million years. H. erectus seems a likely candidate. Whatever the name we choose to give this mystery lineage, what these results show is that gene flow was possible not just among contemporaries but also between ancient and more modern lineages.
Just to show how little we really know of the human story, another genetic surprise has confounded palaeoanthropologists. Scientists succeeded in extracting the most ancient mitochondrial DNA so far, from the Sima de los Huesos site in Atapuerca, Spain.
The morphology of these well-known Middle Pleistocene (approximately 400,000 years old) fossils have long been thought to represent a lineage leading to the Neanderthals.
When the results came in they were actually closer to the 40,000 year-old Denisovans from Siberia. We can speculate on the result but others have offered enough alternatives for me to not to have to add to them.
The conclusion that I derive takes me back to Dmanisi: We have built a picture of our evolution based on the morphology of fossils and it was wrong.
We just cannot place so much taxonomic weight on a handful of skulls when we know how plastic - or easily changeable - skull shape is in humans. And our paradigms must also change.
Some time ago we replaced a linear view of our evolution by one represented by a branching tree. It is now time to replace it with that of an interwoven plexus of genetic lineages that branch out and fuse once again with the passage of time.
This means, of course, that we must abandon, once and for all, views of modern human superiority over archaic (ancient) humans. The terms "archaic" and "modern" lose all meaning as do concepts of modern human replacement of all other lineages.
It also releases us from the deep-rooted shackles that have sought to link human evolution with stone tool-making technological stages - the Stone Ages - even when we have known that these have overlapped with each other for half-a-million years in some instances.
The world of our biological and cultural evolution was far too fluid for us to constrain it into a few stages linked by transitions.
The challenge must now be to try and learn as much as we can of the detail. We have to flesh out the genetic information and this is where archaeology comes into the picture. We may never know how the Denisovans earned a living, after all we have mere fragments of their anatomy at our disposal, let alone other populations that we may not even be aware of.
What we can do is try to understand the spectrum of potential responses of human populations to different environmental conditions and how culture has intervened in these relationships. The Neanderthals will be central to our understanding of the possibilities because they have been so well studied.
A recent paper, for example, supports the view that Neanderthals at La Chapelle-aux-Saints in France intentionally buried their dead which contrasts with reports of cannibalistic behaviour not far away at El Sidron in northern Spain.
Here we have two very different behavioural patterns within Neanderthals. Similarly, modern humans in south-western Europe painted in cave walls for a limited period but many contemporaries did not. Some Neanderthals did it in a completely different way it seems, by selecting raptor feathers of particular colours. Rather than focus on differences between modern humans and Neanderthals, what the examples show is the range of possibilities open to humans (Neanderthals included) in different circumstances.
The future of human origins research will need to focus along three axes:
Sites in the latter category are few and far between. In Europe at least, many were excavated during the last century but there are some outstanding examples remaining. Gorham's and Vanguard Caves in Gibraltar, where I work, are among those because they span over 100,000 years of occupation and are veritable repositories of data.
There is another dimension to this story. It seems that the global community is coming round to recognising the value of key sites that document human evolution.
In 2012, the caves on Mount Carmel were inscribed on the Unesco World Heritage List and the UK Government will be putting Gorham's and associated caves on the Rock of Gibraltar forward for similar status in January 2015. It is recognition of the value of these caves as archives of the way of life and the environments of people long gone but who are very much a part of our story.
Prof Clive Finlayson is director of the Gibraltar Museum and author of the book The Improbable Primate.


Deep-sea worms are not the most exciting or active creatures from looking at them.

But scientists researching them have found that they have one big thing going for them.
The team thinks that these wave dwelling worms could have life spans of up to 300 years or even longer in the deepest parts of our oceans.
Jeanne Calment of France, the oldest human on record, only managed to get to 122 years old!
One problem the team had was, working out how old the creatures could grow to...
So lead researcher, Alanna Durkin from Temple University in Philadelphia, Pennsylvania and her team used information from another study of worms to make a computer program to work it out.
The team think that, Escarpia laminata could be one of the longest living creatures in our oceans!
Alanna also said "It's possible that new record-breaking lifespans will be discovered in the deep sea, since we are finding new species and new habitats almost every time we send down a submersible,"


The Russian Republic of Dagestan, which translates as "land of the mountains", is situated in Russia's turbulent North Caucasus with Chechnya and Georgia to the west, Azerbaijan to the south and the Caspian Sea to the east.

So high are its peaks in some places that certain areas are accessible only by helicopter. The republic is also famed for its ethnic and linguistic diversity, being home to more than 30 languages.
Several dozen Muslim peoples have settled among the high valleys over the centuries.
The Avars form the largest ethnic group and account for about a fifth of the population. A further substantial proportion is made up of Dargins, Kumyks and Lezgins. About 10 per cent are ethnic Russians. There are also Laks, Tabasarans and Nogai, to name but a few of the other significant groups.
Profiles compiled by BBC Monitoring
The republic's constitution declares the protection of the interests of all of Dagestan's peoples to be a fundamental principle. It is a delicate balance to maintain, in what is Russia's most ethnically diverse province.
The republic has oil and gas reserves and also the fisheries potential offered by a share in the resources of the Caspian sea. However, it is prey to organized crime and regional instability. The crime barons may prosper but the people are amongst the poorest in Russia.
Dagestan was the birth place of Imam Shamil, the legendary fighter who in the 19th century spearheaded fierce resistance by tribesmen of Chechnya and Dagestan to the spread of the Russian empire. His name is still revered by many in both republics.
When the Bolsheviks sought to enforce control in the Caucasus in the early 1920s, Dagestan became an autonomous Soviet republic within the Russian Federation. During the Stalinist period, its peoples escaped the mass deportation inflicted on their Chechen neighbours and many others.
After the fall of the Soviet Union, the republic's authorities stayed loyal to Russia, but the region became infamous for its lawlessness and corruption. Organised crime is reported to flourish and kidnappings and violence are commonplace. Firearms are ubiquitous and assassinations are a regular event.
Moscow blames much of this on Chechen-based separatism, but others say lust for profit, combined with a gun culture, is the root cause.
In the 1990s, separatist warlords from neighbouring Chechen openly led armed operations in Dagestan on several occasions. In 1995 and 1996, they seized hundreds of hostages in hospitals in the Dagestani towns of Budennovsk and Kizlyar. Scores died in the attacks.
Dagestan's Muslims, who tend to follow sufism combined with local tradition, generally steered clear of Chechen-style separatism, but after the late 1990s, radical and militant elements said to be linked with the more fundamentalist wahhabist tendency began to gain in influence.
In August 1999, an Islamic body declared an independent state in parts of Dagestan and Chechnya, and called on Muslims to take up arms against Russia in a holy war.
Chechen fighters crossed into Dagestan in support, but within a few weeks, Russian forces had suppressed the insurrection.
The republic has seen numerous bombings targeted at the Russian military stationed in the republic.
Russia accused Dagestani militant Magomed Vagabov of being behind an attack on the Moscow metro by two female suicide bombers from Dagestan in March 2010, in which 39 people died. In August 2010, Russian forces killed Vagabov in Dagestan.


The North East's UKIP MEP Jonathan Arnott has announced his candidacy for the leadership of the party.

The 35-year-old was elected to the European Parliament in 2014 and is a former general secretary of the party.
The leadership race was sparked by the resignation of Nigel Farage, who quit following the EU referendum vote.
Mr Arnott said he wanted to be a "strong and credible voice for the North East".
Many of UKIP's best-known faces have been blocked from contesting the leadership after the Euro sceptic party's ruling National Executive Committee (NEC) said all candidates must have been a member for at least five years.
It means UKIP's only MP, Douglas Carswell, ex-MP Mark Reckless and prominent former spokeswoman Suzanne Evans, who is temporarily suspended from the party, will not be able to enter the contest.
Mr Arnott, said he wanted to appeal to the millions of voters who did not support UKIP in last year's general election, but backed Brexit in the EU referendum.
He warned that the elevation of Theresa May to the premiership meant that "negotiations for the UK's withdrawal from the EU will be in the hands of those who do not actually want us to leave".
He added: "There are many challenges that lie ahead but first we must hold the government's feet to the fire to enact the will of the 17.4 million people who voted for Brexit in the referendum."
Candidates must pay a Â£5,000 deposit to enter the contest, which will be refunded if they secure at least 5% of the vote.
Nominations, which need to be backed by a proposer and 50 supporters from at least 10 branches, close on 31 July.


More than 1,000 people had to be evacuated from the site of a music festival because of storms last night.

The Tiree Music festival-goers were sheltered in schools, the island airport and people's homes.
Organisers said the festival, headlined by The Fratellis and due to run until Sunday night, would get back under way again on Saturday.
Director Daniel Gillespie said: "We've got 1,000 campers under shelter and to do it so quickly has been amazing."
He added: "It's been pretty humbling seeing the response. We were driving round last night taking food and water to the people and even their spirits, their reaction, was incredible as well."
An official statement from festival organisers said they had "no major issues or concerns but have taken this measure as a caution".
It added: "Working closely with Argyll and Bute council, our in-house health and safety team and emergency services, coaches and buses have been arranged for attendees and we have provided accommodation for campers in local schools, business centres and churches.
"We would like to take this opportunity to thank the wonderful people of Tiree who have been tremendous in opening their doors to our visitors."
The festival, founded in 2010, has previously won awards including Best Small Event at the UK Event Awards and Best Small Festival in Scotland at the Scottish Event Awards.
This weekend's line-up includes The Fratellis, Funbox, We Banjo 3, Skerryvore, Manran, Hugh Fearnley-Whittingstall, Gentlemen of Few, Skipinnish, The Chaplins, Cherry Grove, Trail West, Gunna Sound, Dun Mor, The Lowground and Chunks.


A man has been charged over the death of a man who collapsed in the road after being stabbed.

Michael Ward, 40, collapsed in Bush Avenue in Ramsgate at about 17:00 BST on Sunday and died at the scene.
He had been stabbed in the leg in St John's Crescent before he drove to Bush Avenue, Kent Police said.
A 21-year-old man has been charged with murder and is due to appear at Medway Magistrates' Court later.


The UK needs to start fracking to establish the economic impact of shale gas, an industry-funded body has said.

The Task Force on Shale Gas says only after fracking has begun will it be possible to determine how much gas can be recovered.
It said shale could create thousands of jobs and improve UK energy security, but doubted if it would cut prices.
However, the report was criticised by Greenpeace, coming just days after a climate deal was agreed in Paris.
"Whatever planet the UK fracking lobby inhabits, it can't be the same one where world leaders just reached a historic deal that puts fossil fuels on the wrong side of history," Greenpeace said.
"If the UK government is really committed to keeping its end of the Paris deal, it must rethink its support for fracking and back safe, cheap clean energy instead."
The Task Force on Shale Gas said without reliable estimates of how much gas could be recovered, companies would not start to develop the industry.
The report calls on the government and local communities to allow initial exploratory wells.
Chairman of the task force, Lord Smith, said that shale could help cut emissions by replacing more polluting coal: "We need to be even clearer... about shale gas providing a bridge to a low carbon future."
However, he said he doubted whether shale gas would cut UK energy prices "because its almost impossible to tell if that is true or not".
Unlike previous reports, it does not attempt to estimate how much employment the industry could generate, but argued that thousands of jobs could result.
It admitted that the impact on prices of properties affected by fracking was uncertain and that owners of homes near fracking sites could see values fall.
Lord Smith said: "They are understandably worried. The evidence that we can see particularly from the States is that there is a dip in value but then it recovers."
He said that both the industry and government must ensure that any community payment scheme directly compensates property owners: "The people most directly affected should get some direct benefit."
Shale gas production has reduced energy prices in the US, but the UK is part of a wider EU gas market and the volumes of shale gas produced here would be modest in EU terms.
Lord Smith criticised the government's recent decision to axe funding for carbon capture and storage (CCS) technology because it was crucial to creating a shale gas industry.
"One of the things that we highlight in this report is our disappointment that the government's decision to abandon the Carbon Capture and Storage fund," he said.
The report is the fourth and final document from the Task Force on Shale Gas.
The body, which is funded by the fracking industry, was set up "to provide an impartial, transparent and evidence-based assessment of the potential benefits and risks of shale gas extraction to the UK".
An earlier report called for a new regulator for the industry to improve public confidence in the sector.
In July the task force said that fracking could be done safely in the UK with "rigorous regulation".
The government has said it will make a final decision on whether to allow shale gas drilling at two sites in Lancashire, after it was rejected by Lancashire County Council in June. An appeal will be heard next year on plans by the energy firm Cuadrilla to frack at Roseacre and Little Plumpton.


Researchers at Aberystwyth University are attempting to solve the problem of bad tasting beer.

Yeast often suffers heat damage during brewing which can cause a bad taste.
The researchers are trying to understand why some yeast cells survive high temperatures and others do not.
Dr Hazel Davey said: "Our lab strain of yeast recovers well if provided with nutrients during the hot conditions, whereas the brewers' yeast shows better recovery in the absence of nutrients."


Tyrone Gaelic football manager Mickey Harte has paid tribute to his "beautiful" daughter who was murdered.

"She was a wonderful daughter, a brilliant sister for the boys and we will always treasure her," he said.
Michaela McAreavey, 27, was found dead in her bathtub at the four-star Legends Hotel in Mauritius. A post-mortem has revealed she was strangled.
She married John McAreavey, a member of Down's Gaelic football team, on 30 December. The couple were on honeymoon.
Ten people have been questioned over the death. A number of people are still being questioned. Mr McAreavey is not a suspect.
Local reports suggest police are focussing on two suspects.
Police are examining CCTV footage from the complex.
Mr Harte described his daughter as a "gem".
"What a day she had on her wedding day. She was just radiant - a beautiful girl, I just love her to bits and so does all our family.
"Michaela was a wonderful lovely wee girl who grew into a beautiful lady. Just radiant. She is just so special.
"She will always be special to me and this family. We will miss her so much but we love her to bits and her spirit is with us still."
Mr Harte also made a special appeal on behalf of Michaela's husband, John widowed after less than two weeks of marriage.
"We are so sorry. We are equally sorry for poor John whom she adored and who adored her.
"They did not get long to share their lives with each other. It is just such a shame, our hearts are broken.
"John is isolated out there, he has had an awful time. Our hearts go out to him," he said.
"I would ask anybody in the press out there to please respect his privacy."
Mr Harte said he was also speaking on behalf of Mr McAreavey.
"He is devastated, leave him alone. He is in a lonely place. Even his own family members are not out there yet so please back off."
An official from the Irish Embassy in Pretoria, South Africa, is travelling to Mauritius to help Mr McAreavey.
Mr Harte also appealed for privacy for the families and friends to allow them to grieve.
"I know there are lots of people who have had this experience before and we have tried to empathise with them, but you can't get the feeling unless you've been there.
"God save anybody from having to go to this place.
"This is our day to bear this cross."
Police Inspector Ranjit Jokhoo told the BBC that Mr and Mrs McAreavey had been having a cup of tea at an outdoor cafe after using the swimming pool at the Indian Ocean resort.
Ms McAreavey then went back to her room, which was about 50m away, to fetch something, said to be biscuits for her tea, Insp Jokhoo said.
When she failed to return after 20 to 30 minutes, her husband went to the room to look for her. He found her dead in the bathtub in the room.
The inspector said that the killer had used an electronic key card to enter the room, as there had been no forced entry. He said that nothing had been found to have been stolen.


A Cardiff landlord has been fined Â£4,000 after admitting renting five properties to students which inspectors found had "dangerous failings".

Spiridian Micallef from Sully Road in Penarth guilty plea to eight charges relating to a rented property in Albany Road.
Council officials were called in following complaints made last November, city magistrates were told.
Inspectors found faulty electrical sockets and several other issues.


A 30-year-old man has died and two others are missing after a small fishing boat overturned in the Firth of Forth.

Birrell Stewart was picked up from the water about a mile out to sea off the Fife coast at about 17:00 on Monday.
Mr Stewart, from East Wemyss, was declared dead a short time later.
Jason Buchan, 35, and David Stead, 47, both from East Wemyss, who were in the boat fishing for mackerel with Mr Stewart, are missing.
The three amateur fishermen were reported missing amid concerns of worsening weather conditions.
The coastguard launched a search after it was alerted by someone on the shore.
They found the 10ft boat and later located Mr Stewart, who was airlifted to Ninewells Hospital in Dundee, where he was declared dead.
Local councillor Tom Adams, who knows the men, said they were good friends who lived in the East Wemyss area and made regular trips out on the boat fishing for mackerel. He said the tragedy was ''devastating for the whole coast''.
He added that the men enjoyed fishing as a hobby.
Lifeboats from Anstruther and Kinghorn were involved in the search on Monday night, as well as the Rescue 131 RAF helicopter from Boulmer and three coastguard teams from Kinghorn, Leven and St Andrews.
Mark Rodaway, a national maritime operations manager with the Maritime and Coastguard Agency, told BBC Scotland three teams were taking part in a resumed coastline search and a further two RNLI lifeboats had put to sea to re-cover the area in daylight.
He said: "Half an hour into yesterday's search we located the overturned hull of the small fishing boat which gave us extremely good data on the position and 45 minutes after that we recovered the gentleman who, sadly, was subsequently declared deceased.
"So we know where to look. The search area itself remains quite tight and I have got an awful lot of assets to cover that."
Mr Rodaway said the weather was "moderate" at the time of the incident. He said there were fresh breezes but definitely not gale force.
Police said officers were in contact with the families of the missing men.
Jacqueline Thomson, deacon in the Church of Scotland parish of Buckhaven and Wemyss, said: "The community is deeply shocked and saddened by this fishing tragedy, which started off as something that many people in this part of the Fife Coast enjoy doing as a recreational past-time.
"On behalf of the church we would wish to convey that our thoughts and prayers are with the families as they await news of their loved ones."
Matthew Docherty, of East Wemyss angling club who was out fishing on the same day, said: "Conditions were excellent, it was flat like a pancake, but as soon as it got later it started picking up and just rained, it was torrential.
"There was just a big wall coming towards us, so I said lets get back in. We got in and that's when I got the phonecall about them."
Ch Insp Adrian Annandale, of Police Scotland, said: "We have been working closely with colleagues from the coastguard as well as volunteer lifeboat personnel and I would like to thank our partners for all of their assistance and support.
"My thoughts are with the friends and family of all of the fishermen, and with the local community at this time. We will look to provide further updates on our ongoing inquiry once we have more information."


A historic Irish building, designed by the architect of the White House, has been damaged by fire for the second night in a row.

Dublin Fire Brigade was called to Belcamp House in the north of the city in the early hours of Friday and again overnight into Saturday.
The Georgian mansion was designed by Kilkenny native James Hoban - architect of the White House in Washington DC.
Belcamp has been significantly damaged by fire and vandalism in recent years.
The site, off the Malahide Road, was formally used as a secondary school - Belcamp College - but was sold for development in 2004.
Dublin Fire Brigade deployed five units to the site early on Friday morning and a further three units overnight into Saturday.
A spokesman said it was too early to say what had caused this week's fires.
An Taisce (the National Trust for Ireland) had previously included Belcamp on its list of buildings at risk and it assessed it to be under "critical" threat.
Following the first fire on Friday morning, An Taisce wrote to Fingal Council urging the local authority to "take urgent action to protect the 18th Century house".
"This morning's fire has caused serious damage to the ornate plasterwork decorated interiors, which are the finest of the late 18th century period in Fingal.
"The extent of damage is still undetermined," the trust said.
An Taisce claimed the building had been "inadequately secured" following the Irish property crash.
It called on the council to "urgently intervene to establish the current ownership of the building and any continuing involvement of NAMA (National Assets Management Agency) and to ensure that the roof is repaired and further damage averted".


Jessica Ennis says she will not run in the 100m hurdles following her Olympic triumph in the heptathlon.

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The 26-year-old from Sheffield considered competing in the event, which starts on Monday, after running a personal best time of 12.54 seconds in the first discipline of the heptathlon.
"I did think about it, especially when I  she said.
"But for me it was just about the heptathlon. I'm more than happy with that and I just need to rest now."
Not surprised Jess isn't doing hurdles. These next few days & weeks are dreamlike and she needs to capture every second.
Olympic gold medallist Denise Lewis on Twitter
Ennis, who missed out on competing in the Beijing Olympics because of injury, clocked a time in the first event of the heptathlon that would have won gold at the 2008 Games. It was also a world best in the multi-event discipline.
But the Sheffield athlete told BBC Breakfast she wanted to rest and revel in becoming Olympic champion for the first time.
"Right now it is just to enjoy this moment for as long as I can," she said. "I want to spend some time with my family and then just have a bit of a break. It has been such a long build-up and stressful. I am looking forward to relaxing."
Ennis's gold came on a sparkling Saturday night for British athletics in the space of an hour at the Olympic Stadium.
Following her victory, Greg Rutherford won a surprise gold in the men's long jump before Mo Farah stormed to victory in the men's 10,000m.
Having been billed as the 'face' of the Games, Ennis said she was relieved to have won in a home Olympics and said she kept the medal by the side of her bed overnight.
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"I slept for about two hours," she told BBC Radio 5 live. "I just kept looking at my medal and thinking about what I've achieved. I don't think it's sunk in yet. I looked at it as soon as I woke up and I'll look at it every day.
"There has been lots and lots of pressure and everyone just expected me to win. It's been quite hard to deal with that but I've had great people around me.
"I just wanted to get focused and use the positivity from everyone around the country and use that to get this gold medal."
On her plans for the immediate future, she said: "I'm definitely going to relax, eat lots of rubbish food, have a few glasses of wine and enjoy this moment for as long as possible."


Exeter are set to sign a lock from a Pro12 club and an overseas back.

The Premiership side have been hit by injuries to key players in both positions in recent months.
Chiefs are also close to signing a second lock who will join the club in the summer and extending the contracts of a number of the existing squad.
"We've been pretty active over the last two or three weeks, we're pretty satisfied to where we're going to be this year," head coach Rob Baxter said.
Mitch Lees and Damien Welch have both been ruled out for much of the remainder of the season, leaving England second-row Geoff Parling as the only experienced lock available to the Chiefs.
While Baxter would not reveal who he was in talks with, he said he hoped to announce the signing in the next few days.
"He's a player invoked in the Pro12 at the moment. We think it's a really good fit for us and people will understand that when they see the player it is," said Baxter.
"Initially with the first injury to Damien we were thinking we could get through things and we had enough cover, but the second injury to Mitch really tipped the balance too far for us to try and battle our way through.
"We've still got another Champions Cup game this weekend and then we've got 10 Premiership games on the bounce."
Exeter currently have England centre Henry Slade out for much of the rest of the season, while Sam Hill, Michele Campagnaro and Jack Nowell have all been called up for Six Nations duty, meaning Baxter is also short on backs.
"We've still got to wrap up quite a few details on visas etc," said Baxter regarding the overseas signing.
"Hopefully by mid-February we'll have someone over here and that will then complete the look of our backs.
"Bringing in some extra cover allows us to increase our flexibility. Having guys who can cover one or two or three positions will be very important for us in this next 10-week block."


(Closed): US stocks closed higher in November's first trading session, boosted by positive US construction data.

The Dow Jones Industrial Average was up 0.94% at 17,828.76 points, while the Nasdaq was 1.45% higher at 5,127.15 points.
At the same time, the S&P 500 index was up 1.19% at 2,104.05 points.
Data from the Commerce Department showed construction of new homes and highways reached a seven-year high.
Petroleum companies had a strong day. ExxonMobil was up 3.1% and Chevron was up 4.5%.
Credit card firm Visa failed to impress investors with its plan to buy Visa Europe, currently under separate ownership.
The firm, which also announced lower than expected quarterly profits, topped the losers' list on the Dow, falling 3%.
Valeant Pharmaceuticals was up 7.2% after weeks on a downward streak.
The drug maker was accused by short seller Citron, of using a speciality pharmacy to misprice drugs. Shares rebounded on the news Citron would not publish new allegations.


Arsenal and England midfielder Jack Wilshere scored his first goal for 10 months as the Gunners' Under-21 team lost 2-1 to West Brom on Monday.

The 24-year-old made his return against Newcastle Under-21s earlier this month after breaking his fibula in August.
Before Monday, his last goals were for his country as Roy Hodgson's side beat Slovenia 3-2 in June.
Wilshere featured in five of England's first six qualifying fixtures for this summer's European Championship.


A construction worker who died following an incident on the new Queensferry Crossing has been named as John Grant Cousin.

The 62-year-old from Northumberland was working on the Â£1.4bn bridge which is being built across the Firth of Forth.
He is thought to have been hit by a moving boom on a crane on the deck of the north tower on 29 April.
An investigation is being carried out by Police Scotland and the Health and Safety Executive.
In a statement, Police Scotland said Mr Cousin's family wanted to thank everyone for their "thoughts and support at this sad time".
It is understood that he suffered severe blood loss in the incident and he was unable to be resuscitated.
Another man was injured in the incident.
The bridge, between Fife and Edinburgh, is due to be completed at the end of the year.
Work started in 2011 and Mr Cousin's death is thought to be the first fatality at the site.


A man who has lived with a stammer since the age of three has graduated after qualifying as a speech and language therapist.

Richards Stephens, 37, from Bury, said he could not say his own name as a child and "wouldn't speak up in class".
He decided to train as a speech therapist after his confidence was boosted through his work as a psychiatric nurse and a support group.
He graduated at the University of Manchester with first-class honours.
"Instead of going out at night, I've been reading journals," he told BBC Radio Manchester.
"Now I have a mild stammer but as a child it was very strong. I could never be fluent in a sentence so it was on your mind constantly.
"You fear talking, you fear being put in a social situation - it's a struggle because you know the words are there, it's just they won't come out for whatever reason," Mr Stephens said.
Treated by a speech and language therapist from the age of seven, he said therapy failed to deal with his psychological and social problems.
"Stammering used to dominate all of my life choices. I first chose to study IT as an undergraduate at Liverpool John Moores because I thought I wouldn't have to talk much and could communicate via e-mail.
"In restaurants I'd order dishes which were easier for me to pronounce and not the ones I really wanted."
He gained confidence in his early 30s after attending a support group set up by the British Stammering Association and said his voluntary work at American summer camps for children who stammer had been a "revelation".
"I want to help better understand this speech disorder, not only for myself, but for the benefit of the millions of people who stammer worldwide.
"The stammer is only a small part of what we train in because we also train in how to work with people who have had strokes, language problems and delays."