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Four senior aides to Boris Johnson have resigned from Downing Street within hours of each other amid growing pressure on the prime minister.
Policy chief Munira Mirza left over what she said was a "scurrilous" attack by the PM on Labour's Sir Keir Starmer.
Three staff caught up in the lockdown parties row - including Martin Reynolds, who sent out a "bring your own booze" invitation - also quit.
It came as the PM faces more questions over his leadership from his party.
Director of communications Jack Doyle confirmed his exit shortly after the departure of Ms Mirza.
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.
A statement from a No 10 spokeswoman said chief of staff Dan Rosenfield had offered his resignation to the prime minister earlier on Thursday, but would stay on while his successor was found.
And Mr Reynolds - the prime minister's principal private secretary - will do the same, but then return to a role at the Foreign Office.
A number of MPs supportive of the prime minister have been tweeting praise, suggesting Mr Johnson was responsible for making necessary staff changes following a damning report by civil servant Sue Gray into rule-breaking parties in No 10 during the pandemic.
Energy Minister Greg Hands told BBC Breakfast the resignations came after Mr Johnson "made it clear there would be a shake-up" at Downing Street.
Speaking to BBC Radio 4's Today programme, Conservative backbencher Huw Merriman said many MPs were being loyal to the PM and concentrating on the positives. But he said he was "deeply troubled" by the situation and agreed that the prime minister should "shape up or ship out".
Ms Mirza quit over the PM's false claim that Sir Keir 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."
Labour's deputy leader Angela Rayner said: "With [Mr Johnson's] senior advisers and aides quitting, perhaps it is finally time for him to look in the mirror and consider if he might just be the problem."
The chief of staff, the chief of communications, the official who runs the prime minister's private office, and his policy chief - all gone. A quartet of departures from Number 10.
In the words of one senior backbencher, not just a nightmare on Downing Street, but a total meltdown.
Yet two different things happened on a turbulent day. The first three paid the price for the fiasco over lockdown gatherings - one sending the invite to the garden drinks, the other two crafting Boris Johnson's bungled response to weeks of damaging claims.
Boris Johnson's backers claim it's evidence that he'll keep his vow - 'I get it, I will fix it' - that Number 10 can become the home to a new and improved operation.
The resignation of Munira Mirza is of a very different order. For more than 10 years she's been one of Boris Johnson's true political friends - described as 'Boris' brain' by one former colleague, an ally of ideas.
The resignations come as backbench unrest is growing within the Conservative Party.
The BBC is aware of 17 Tory MPs who have submitted letters of no confidence in the prime minister - 54 are needed to trigger a leadership contest.
Many have cited the PM's participation in parties with staff in No 10 during lockdowns as their motivation to challenge Mr Johnson.
Who are the aides who resigned?
The head of policy at No 10, Munira Mirza worked for Boris Johnson for 14 years, including when he was mayor of London, and was seen as one of his most trusted advisers.
Jack Doyle worked as a Daily Mail journalist before joining Downing Street. He is reported to have attended one of the Christmas gatherings in December 2020, where he thanked staff for their work.
Martin Reynolds is a civil servant and worked as Boris Johnson's Principal Private Secretary. In that role, he headed up the PM's private office. He is responsible for sending an email invitation to staff inviting them to socially distanced drinks in the No 10 garden in May 2020.
Dan Rosenfield joined Number 10 as chief of staff in January 2021, and had worked in the Treasury for former chancellors Alistair Darling and George Osborne.
Other MPs have pointed to the PM's remark about Sir Keir as one of their reasons for writing a letter.
In a debate on the findings of a report into lockdown parties 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.
Savile was a serial sexual abuser who attacked hundreds of women and children at 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.
Speaking to Channel 5 News after Ms Mirza quit - but before the news of the other resignations was made public - the prime minister said he was "sorry to lose Munira".
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."
Downing Street confirmed that Tory MP Andrew Griffith would take over from Ms Mirza as the head of the PM's policy unit, as well as become the minister for policy.
Speaking to BBC Newsnight, ex-Tory foreign secretary Malcolm Rifkind criticised Mr Johnson's "very hollow" political judgement and described the situation as "not so much the end of the beginning as the beginning of the end".
These gatherings included a "bring your own booze" garden party in May 2020 - the email invite to which was sent by Mr Reynolds.
The Metropolitan Police has launched an investigation into 12 of the parties and a full report by Ms Gray is expected to be published afterwards.
