Four senior aides to Boris Johnson have resigned from Downing Street within hours of each other amid growing pressure on the prime minister.
Director of communications Jack Doyle confirmed his exit shortly after the departure of policy head Munira Mirza.
They were followed by the chief of staff Dan Rosenfield and senior civil servant Martin Reynolds.
The top aides’ resignations come as Mr Johnson faces increasing questions over his leadership from within his party.
Mr Doyle told staff that “recent weeks have taken a terrible toll on my family life”, but that he had always intended to leave after two years.
However, Ms Mirza left over the PM’s false claim that Sir Keir Starmer failed to prosecute Jimmy Savile when he was director of public prosecutions, and his refusal to apologise.
Chancellor Rishi Sunak publicly distanced himself from the PM’s original comment, saying: “Being honest, I wouldn’t have said it.”
And asked if Mr Johnson should apologise, he said: “That’s for the prime minister to decide.”
A statement from a No 10 spokeswoman said Mr Rosenfield had offered his resignation to the PM earlier on Thursday, but would stay on while his successor was found.
Mr Reynolds – who was the principal private secretary to the prime minister – will do the same, but then return to a role at the Foreign Office.
The wave of resignations comes at a tumultuous time for Mr Johnson as backbench unrest is growing with the Conservative Party.
The BBC is aware of 17 Tory MPs who have submitted letters of no confidence in the prime minister, with a total of 54 needed to trigger a leadership contest.
Many have cited the PM’s participation in parties at No 10 during Covid lockdowns as their motivation to challenge Mr Johnson.
But some have also pointed to his remark about Sir Keir, including senior backbencher and Defence Committee chair Tobias Ellwood.
On Monday, Mr Johnson accused the Labour leader of spending “most of his time” as director of public prosecutions (DPP) “prosecuting journalists and failing to prosecute Jimmy Savile”, despite having no evidence to back up the claim.
Disc jockey and TV personality Savile was revealed to be a serial sexual abuser after his death in 2011, having attacked hundreds of women and children at various locations including hospitals, schools and the BBC.
By Thursday, Mr Johnson appeared to have backed down, telling reporters he had not been talking about Sir Keir’s “personal record” when he was DPP, adding: “I totally understand that he had nothing to do personally with those decisions.”
But in her resignation letter, published by The Spectator, Ms Mirza said the PM should have apologised for the misleading remarks.
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Speaking to Channel 5 News after Ms Mirza quit – but before the new of all the resignations were made public – the prime minister said he was “sorry to lose Munira”, who had worked with him since he became Mayor of London in 2008.
He said she had “done an outstanding job” and had been “instrumental in delivering all sorts of fantastic policies”.
But asked about her conclusion that his remarks about Sir Keir had been “inappropriate and partisan”, Mr Johnson replied: “Well I don’t agree with that.”
He said: “No-one is commenting, least of all me, about the personal involvement of the leader of the opposition in the handling of that case.
“All that I’ve said is that the leader of the opposition apologised for the CPS’s handling of that issue during his tenure and that’s all frankly that needs to be said on that matter.”