Boris Johnson's premiership has been plunged into fresh crisis after the resignations of four senior Downing Street aides in a single day.
His policy chief Munira Mirza dramatically announced she was quitting over his use of a Jimmy Savile smear against Keir Starmer.
No 10 spin chief Jack Doyle then announced he was resigning and told staff the Downing Street partygate saga had taken a "terrible toll" on his family life.
The PM's principal private secretary Martin Reynolds, who invited aides to a BYOB bash in the Downing Street garden in May 2020, and chief of staff Dan Rosenfield then offered their resignations.
The news will further rock Mr Johnson's premiership as he fights to survive probes by top civil servant Sue Gray and Scotland Yard into lockdown-breaking parties.
Downing Street said Mr Reynolds and Mr Rosenfield had resigned after speaking to Mr Johnson - but speculation was rife that the pair had been sacked.
Asked if it was like the last days of Rome, Treasury minister Simon Clarke replied: "The last days of Rome, I think, were more fun."
To add to the embattled PM's woes, Chancellor Rishi Sunak also issued a public rebuke over his use of the Savile slur against the Labour leader during a Downing Street press conference.
The PM promised MPs he would overhaul the No10 operation after the disastrous flood of allegations of rule-breaking parties in No 10 and Whitehall during the pandemic.
But Mr Johnson, who had been hoping his premiership had stabilised, is now once again fighting for his political life.
A Downing Street source said: "There’s meant to be an all-staff meeting tomorrow. Assuming there are still any staff."
The departure of Ms Mirza, a longtime ally who worked with Mr Johnson when he was London mayor, was unexpected - and not linked to the partygate row.
Her anger centred on his failure to give a full apology for trying to claim Mr Starmer had failed to prosecute Jimmy Savile in 2009 when he led the Crown Prosecution Service - something the CPS later apologised for.
But independent fact-checkers Full Fact found it had never been suggested that Mr Starmer was personally involved in the decision.
Mr Johnson initially doubled down but finally admitted today that Mr Starmer "had nothing to do personally with those decisions".
"I want to be very clear about this because a lot of people have got very hot under the collar," Mr Johnson told reporters.
"I'm talking not about the leader of the opposition's personal record when he was when he was DPP and I totally understand that he had nothing to do personally with those decisions."
But his failure to fully apologise angered his long-time aide.
"I believe it was wrong for you to imply this week that Keir Starmer was personally responsible for allowing Jimmy Savile to escape justice," Ms Mirza said in a letter seen by The Spectator.
"There was no fair or reasonable basis for that assertion. This was not the usual cut and thrust of politics; it was an inappropriate and partisan reference to a horrendous case of child sex abuse.
"You tried to clarify your position today but, despite my urging, you did not apologise for the misleading impression you gave.
"You are a better man than many of your detractors will ever understand which is why it is so desperately sad that you let yourself down by making a scurrilous accusation against the Leader of the Opposition."
Mr Johnson, who once described her a "brilliant thinker" and listed her as one of the five women who had influenced and inspired him the most, said he was "sorry" to lose her.
Ex-No 10 aide Dominic Cummings tweeted praise for Ms Mirza's "moral courage", and added: "It's also an unmistakeable signal the bunker is collapsing & *this PM is finished*."
Soon after, Mr Doyle, the Downing Street director of communications, told staff he was leaving but insisted his intention was always to leave after two years.
According to the Daily Mail, he told aides: 'It was always my intention to do two years. '
"Recent weeks have taken a terrible toll on my family life.
"Thank you all for your hard work and dedication. I wish you all the best for the future."
A No 10 spokesperson said: "Jack Doyle has left government.
"He has made a huge contribution and the Prime Minister is immensely grateful for the work he has done."
Mr Doyle, a former Daily Mail journalist, was appointed as Director of Communications in April 2021 after becoming Boris Johnson's press secretary in 2020.
He is believed to have attended at least two of the parties in Downing Street and Whitehall being investigated by Sue Gray.
No 10 then confirmed the departures of Dan Rosenfield and Martin Reynolds, who had both been in the firing line over the partygate row.
A spokesperson said: “They will continue in their roles while successors are appointed, and recruitment for both posts is underway."
Meanwhile, Tory Mayor Andy Street said Mr Johnson's behaviour over partygate was "bad by any measure".
In comments that will alarm Downing Street, the influential West Midlands Mayor told BirminghamLive: "What he's done, very clearly, is wrong. It's bad by any measure.
"And I find it very hard to understand how on earth it could have happened."
But he stopped short of calling for him to go - and said he believed Mr Johnson was sincere in his apology.
