Western and UN leaders have condemned a daylight Russian missile barrage that hit Ukraine’s largest children’s hospital, leaving an unknown number trapped under the rubble, as strikes across the country killed 36 people in one of the deadliest attacks this year.

Volodymyr Zelenskiy vowed retaliation as he said Kyiv’s Okhmatdyt hospital, the main treatment centre in the country for children with cancer, had taken a direct missile hit. The strike was part of one of the heaviest attacks on the capital since Russia’s February 2022 invasion.

“Russia cannot help but know where its missiles are flying, and must fully answer for all its crimes: against people, against children, against humanity in general,” Zelenskiy posted on the Telegram messaging app.

A spokesperson for António Guterres, the UN secretary general, said he strongly condemned the “particularly shocking” strikes against the children’s hospital and another medical facility, and the security council is to meet on Tuesday at the request of Britain, France, Ecuador, Slovenia and the US.

Barbara Woodward, the UK ambassador to the UN, said on X: “We will call out Russia’s cowardly and depraved attack on the hospital.” The Italian foreign minister, Antonio Tajani, called the missile strike “war crimes”.

The British prime minister, Keir Starmer, condemned “attacking innocent children” as the “most depraved of actions”.

The strikes across Ukraine on Monday highlighted the country’s air defence vulnerabilities a day before Nato leaders are set to meet in Washington for their annual summit, where they are expected to announce new measures to aid Ukraine. Twenty-two people were killed in the capital, including two staff members at the hospital, where three children were hurt.

The strike largely destroyed the hospital’s toxicology ward, where children with severe kidney issues were being treated. Hundreds of rescue workers and volunteers joined the effort to clear the debris and search for survivors. Officials and emergency staff said it was not immediately clear how many doctors and patients – dead or alive – remained trapped under the rubble.

“We are extracting whoever we can. We don’t know the number of people trapped there,” said the health minister, Viktor Liashko, outside the hospital.

Images from inside the hospital, which treats 20,000 children annually, showed bloodied children, collapsed ceilings and destroyed operating rooms.

Maria Soloshenko, 21, a nurse in the toxicology ward, said hospital staff had been in the process of moving the children to a bomb shelter when the explosion occurred. “There was immediate panic when the strike hit,” she said, her gloves covered in blood.

Soloshenko said children as young as 18 months had to be urgently taken off dialysis and quickly evacuated through the building’s windows.

The Guardian witnessed many young cancer patients in distress during the evacuation, some barely clothed and with medical tubes still attached to them.

Tanya Lapshina, a nurse in the trauma department, where the facade was ripped off by the blast, said they managed to move all the children to a bomb shelter. She said: “It was complete chaos. The children were in panic, crying in the bunker. There are no words for this. It is awful. I am still shaking.”

Ukraine’s presidential office published an image showing one child with a head injury.

Kyiv’s mayor, Vitali Klitschko, described the attack on the capital as one of the heaviest since Russia’s invasion began more than two years ago. Thanks to western-supplied defences, the city had experienced a relatively peaceful period before Monday.

Russia, which has targeted civilian infrastructure throughout the war, denied responsibility for deaths on Monday. In a statement, the defence ministry attributed the incident, without directly referencing the hospital blast, to Ukrainian anti-aircraft missiles, despite visual evidence that appeared to point to a Russian strike.

Andriy Yermak, the head of the Ukrainian president’s office, shared an image on X that appeared to show a Russian missile over Kyiv moments before it struck a hospital, identifying it as a Kh-101 cruise missile. Ukraine’s Security Service said it found wreckage from the cruise missile, which flies low to avoid detection by radar, at the site.

The Ukrainian prosecutor general, Andriy Kostin, said he had discussed the attack with the international criminal court (ICC) prosecutor, Karim Khan. “We are sending all information and evidence about attacks on Kyiv and other Ukrainian cities to the ICC prosecutor’s office,” he said on national television. The court last month issued arrest warrants for Russia’s former defence minister and its military chief of staff over attacks on Ukraine’s electricity network.

The search efforts at the hospital were hindered by air-raid alarms that forced emergency staff to take shelter.

Shortly after the strike on the children’s hospital, missile debris fell on a separate medical centre in the Dniprovsky district of the city, the mayor said, with seven people thought to be killed.

Strikes were also reported in other parts of the country. In Kryvyi Rih, Zelenskiy’s home town, 11 people were killed and 31 were injured, said Oleksandr Vilkul, the mayor. Another four people died in Pokrovsk in eastern Ukraine when missiles hit an industrial facility, said the Donetsk regional governor.

In total at least 36 people across the country were killed, Ukrainian officials said. Zelenskiy said 40 missiles were used in the attack.

The International Rescue Committee said the attack was part of a broader pattern of Russian strikes on medical facilities since the invasion.

“Since the beginning of the full-scale invasion, the WHO [World Health Organization] has recorded nearly 1,700 attacks on healthcare facilities in Ukraine,” the organisation said in a statement.

During a press conference in Warsaw after a meeting with the Polish prime minister, Donald Tusk, Zelenskiy urged Kyiv’s allies to make a decisive response to the attack.

“I would also like to hear from our partners [about] a greater resilience and a strong response to the blow that Russia has once again dealt to our people, to our land, to our children,” he said, adding that he was waiting for concrete steps from the west to strengthen Ukraine’s air defences and protect its energy sector.

After one of the worst Russian missile strikes against Ukraine in recent months, Nato leaders will sit down in Washington DC this week to announce the details of a hard-fought aid package that will include crucial air defense systems meant to protect Ukrainian cities.

The package put forward by Nato countries has been presented as “historic” and is an widely seen as an attempt to “futureproof” continued aid to Ukraine – but it may not fully satisfy Kyiv, which has been facing unprecedented attacks against civilian sites and infrastructure.

The resumption of large-scale missile strikes against targets in Kyiv will increase the sense of urgency around the discussions among 32 Nato leaders. Images from Kyiv showed children at a pediatric cancer hospital covered in blood and dust after the strike on Monday which a Biden administration official described as “horrific, tragic, senseless”. There were believed to be bodies still trapped under the rubble of the hospital.

“This is a fully deliberate action, specifically designed and approved by … Putin. On the eve of the @Nato summit. As a slap in the face to the alliance,” wrote Mykhailo Podolyak, an adviser to the Ukrainian presidential administration. He called it an “informal signal” that “even the outright murder of children will not make them [the Alliance] take all the necessary decisions. And that is why we continue to attack.”

Observers expect Nato members to pledge of at least four additional Patriot missile batteries to Ukraine at the conclusion of this week’s summit. Zelenskiy had previously asked Nato for seven batteries, telling Nato members that Putin “must be brought down to earth, and our sky must become safe again … And it depends fully on your choice … [the] choice whether we are indeed allies.”

It is expected that the four Patriot missile systems will likely be delivered by the US, Germany, Romania and a Dutch-led multinational effort. Spain, Greece and Poland also field Patriot missile systems but have so far not pledged to supply any batteries to Ukraine. Another system could be provided by Israel, which now employs the Iron Dome and other air defense systems to protect against rocket and missile attacks.

“It’s clear that allies need to step up and provide Ukraine with additional air defense systems, precisely in order to be able to prevent types of tragedies that we’ve seen today, but sadly that we’ve seen time and again, month after a month since the beginning of this brutal and senseless war,” said Michael Carpenter, senior director for Europe at the US National Security Council.

The new military aid package to Ukraine is expected to include a joint commitment from Nato members to spend at least €40bn ($43bn) in 2025 on aid to Ukraine.

“Since Russia’s full-scale invasion in 2022, Allies have provided €40bn in military support to Ukraine each year,” Jens Stoltenberg, the secretary general of Nato, said last month. “We must maintain this level of support as a minimum, and for as long as it takes.”

Stoltenberg had asked for a multi-year pledge from the 32 Nato member states but it did not appear that they had come to an agreement on the eve of the summit.

A European official said that the idea of a multi-year pledge was “still being discussed because some allies including here are uncomfortable with the idea of a multi-annual pledge because of their legal and institutional limitations, so I think we still have to wait”.

One way around that issue, the official said, would be to “just make an annual pledge and then to recommit summit after summit”. Next year’s Nato summit is set to take place at the Hague.

But that could take place after the re-election of Donald Trump, who has threatened to cut aid to Ukraine or make it conditional on starting talks with Russia.

“The big orange elephant in the room for the Nato Summit is that everything good that’s going to be said about Ukraine comes with a big caveat,” said Camille Grand, a former Nato assistant secretary general who is now at the European Council on Foreign Relations. “It is will this all hold if Trump is elected? And I don’t believe in bureaucracies … because it has been agreed at the Nato summit that the Trump administration would follow that.”

Nato members are expected to announce the establishment of a new military command in the German city of Wiesbaden which would coordinate military aid and training for Ukraine, effectively replacing the US-led Ukraine Defense Contact Group. That would in effect move the onus of supplying Ukraine from the Pentagon to Nato, in what US officials have said would be a “bridge to membership” preparing the country to be ready to work with the alliance when it is admitted “on day one”.

The new effort is also seen as a way to “Trump-proof” future aid to Ukraine if he is elected in November by “institutionalising” aid to Kyiv.

Carpenter also said that there would be announcements concerning the provision of F-16s to Ukraine. Belgium, Denmark, the Netherlands and Norway have pledged to provide Ukraine with about 80 US made F-16 fighter jets between them, but the program to get the planes in the air has been hit by delays in delivery and training. The first F-16s are expected to arrive this summer.

On the eve of the summit, diplomats said there was still “no consensus” on Nato issuing an invitation to Ukraine at the summit to join the alliance. “Some allies are reluctant in that direction, but we are discussing languages to at least showcase that Ukraine’s path to membership is irreversible, that there is no there is no way back,” said a European official.

A Biden administration official declined to directly discuss the language of the final communiqué because it was “still being negotiated”, but said that the summit declaration “will include very strong signals of Allied support for Ukraine on its path to Euro-Atlantic integration. And it’s going to also underscore the importance of Ukraine’s vital work on democratic, economic and security reforms.”

Joe Biden came out swinging on Monday against critics of his calamitous June debate performance, telling Democrats in an open letter and Americans in a pugnacious live TV interview he is staying in the presidential race – rejecting growing calls to concede that at 81 he is too ineffective to beat Donald Trump and should drop out in favour of a younger candidate.

The president lashed out at “elites in the party” in a live telephone interview with the MSNBC show Morning Joe, saying they were behind calls for him to quit.

He added: “If any of these guys don’t think I should run, run against me. Go ahead. Announce for president – challenge me at the convention!”

Touting what he said was a demanding campaign schedule around hosting a Nato summit this week, the president insisted: “The American public is not going to move away from me.

“I’m here for two reasons, pal. One, to rebuild the economy for hard-working middle class people, to give everybody a shot. It’s a straight shot. Everybody gets a fair chance. Number two, people always talk about how I don’t have the wide support. Come on, give me a break. Come with me. Watch.”

Concerns about Biden’s age have dogged his time in office but they exploded into open view late last month after the first of two scheduled debates with Trump.

Onstage in Atlanta, Biden appeared hesitant, confused and physically diminished, struggles aides put down to a cold and jet lag.

In comparison, Trump spewed lies virtually unchecked by his opponent or CNN moderators working to rules that precluded instant fact checks.

The result was a polling bump for Trump and panic among Democrats. By Monday, nine House Democrats had called for Biden to quit. A reported move towards a similar call in the Senate did not produce a result.

Biden insisted his poor debate was down to health issues.

“I was feeling so badly before the debate,” he told MSNBC. “They tested me, they thought maybe I had Covid, maybe there was something wrong, an infection or something. They tested me, they gave me those tests. I was clear. So, I had a bad night.”

Touting public appearances since the debate, Biden said he was in vigorous health and out meeting voters more than Trump.

“I have a neurological test every single day sitting behind his desk and making these decisions,” Biden said. “You know it, they know it. I’m not bad at what I do.”

Signaling the size of Biden’s problem, however, the New York Times cited White House visitor logs when it reported that “an expert on Parkinson’s disease” visited “eight times in eight months from last summer through this spring, including at least once for a meeting with President Biden’s physician”.

Andrew Bates, a White House spokesperson, told the paper: “A wide variety of specialists … visit the White House complex to treat the thousands of military personnel who work on the grounds.”

Bates also said Biden had been seen by a neurologist once a year, finding “no sign of Parkinson’s and he is not being treated for it”.

In his open letter to Democrats, the president said he was “firmly committed to staying in this race, to running this race to the end, and to beating Donald Trump”.

He also claimed that in “extensive conversations with the leadership of the party, elected officials, rank and file members and most importantly Democratic voters”, he had “heard the concerns that people have – their good faith fears and worries about what is at stake in this election. I am not blind to them.

“Believe me, I know better than anyone the responsibility and the burden the nominee of our party carries. I carried it in 2020 when the fate of our nation was at stake.”

Biden defeated Trump handily then. But on inauguration day, he was 78 – as old as Trump is now but the oldest man ever to take the presidential oath.

On Monday, Biden said: “I wouldn’t be running again if I did not absolutely believe I was the best person to beat Donald Trump.”

It is unclear what mechanism could be used for replacing Biden, whether with his vice-president, Kamala Harris, or another candidate.

In his letter, Biden pointed to his easy primary win over Dean Phillips, a Minnesota representative who campaigned on the issue of Biden’s age. The president also pointed to the independent Robert F Kennedy Jr, who threatens to take votes in battleground states.

“Do we now just say this process didn’t matter?” Biden asked. “That the voters don’t have a say?

“I decline to do that. I feel a deep obligation to the faith and the trust the voters of the Democratic party have placed in me … it was their decision to make. Not the press, not the pundits, not the big donors, not any selected group of individuals, no matter how well intentioned. The voters – and the voters alone – decide the nominee.

“How can we stand for democracy in our nation if we ignore it in our own party? I cannot do that.

I will not do that.”

Biden said he had “no doubt” he would beat Trump, touting achievements in office. He also said that in a second term, with a Democratic-controlled Congress, he would restore abortion rights by enshrining them in law, while bringing “real supreme court reform” – an ambitious statement, given a Senate map highly favourable to Republicans.

Finally, Biden said he was “standing up for American democracy”.

His letter invoked the 6 January 2021 US Capitol attack carried out by Trump supporters, saying his White House predecessor “has proven that he is unfit to ever hold the office of president. We can never allow him anywhere near that office again. And we never will.

“We have 42 days to the Democratic convention and 119 days to the general election … it is time to come together, move forward as a unified party, and defeat Donald Trump.”

The Michigan governor, Gretchen Whitmer, said she would not run for the Democratic nomination for president this year even if Joe Biden cedes to growing pressure and steps aside.

“It’s a distraction more than anything,” Whitmer told the Associated Press, in an interview to promote her new memoir, True Gretch, which will be published on Tuesday.

“I don’t like seeing my name in articles like that because I’m totally focused on governing and campaigning for the [Biden-Kamala Harris] ticket.”

Whitmer was referring to a Politico report last week, which said that after Biden’s disastrous debate against Donald Trump in Atlanta in June, Whitmer called the president’s re-election campaign chairperson, Jen O’Malley Dillon, to say Michigan – a battleground state – was no longer winnable.

In her immediate response to that report, Whitmer said she was “proud to support Joe Biden as our nominee and I am behind him 100% in the fight to defeat Donald Trump”.

Politico said its source was “someone close to a potential 2028 Whitmer rival for the Democratic presidential nomination”.

Speaking to the AP, Whitmer said: “I think it’s frustrating that there are news outlets that will publish something that a potential future opponent’s staff person would say.”

Few doubt Whitmer will run for the Democratic presidential nomination in 2028. Other governors thought likely to run for that nomination include Wes Moore of Maryland, Andy Beshear of Kentucky, Gavin Newsom of California and Josh Shapiro of Pennsylvania.

Biden is attempting to quash calls for him to quit from those who believe that at 81 he is too old and infirm to campaign and govern effectively. The volume of those concerns amplified considerably after his 27 June debate with his presidential predecessor.

On Monday, Biden lashed out at “elites in the party”, telling MSNBC: “If any of these guys don’t think I should run, run against me. Go ahead. Announce for president – challenge me at the convention!”

Whitmer says she will not do that but her ambitions remain on display.

As the Guardian revealed last week, her book ends with a passage from Theodore Roosevelt’s famous “Man in the Arena” speech, about the need for leaders to take action.

Speaking in Paris in 1910, the 26th president said in part: “The credit belongs to the man who is actually in the arena, whose face is marred by dust and sweat and blood; who strives valiantly; who errs, who comes short again and again, because there is no effort without error and shortcoming.”

In her book, Whitmer adds: “Though these words were written more than a hundred years ago, they’re just as true today – except for two things. The ‘man’ may be a woman. And she may just be wearing fuchsia.”

Emmanuel Macron has asked Gabriel Attal to stay on temporarily as France’s prime minister to maintain stability after a snap general election left the country facing a hung parliament and fraught negotiations to form a new government.

Parties on the left want to seize the moment after their shock win over the far right and on Monday met to discuss policy and potential prime minister candidates. The green-left alliance, the New Popular Front (NFP), surprised pollsters by coming first in the final round – a win that was considered highly unlikely, with pre-election polls predicting a far-right surge into the lead.

However, with no absolute majority, efforts to form a new French government may take weeks.

The snap election was called last month after a humiliating defeat to the far-right National Rally (RN) of Marine Le Pen in European parliamentary elections.

The NFP won 182 seats in the 577-seat national assembly, with Macron’s centrist Together coalition returning 168 deputies and the RN – which after the first round on 30 June had been eyeing a majority – finishing third on 143.

With no single group securing an absolute majority, the options include a technocratic government of experts, the NFP trying to form a minority government and seeking bill-by-bill support, or a broad coalition of the centre left and centre right.

Attal tendered his resignation on Monday morning after Macron’s camp lost more than a third of its MPs. The president asked Attal to remain in power in a caretaker capacity to see out the period of the Paris Olympics and to reassure the international community and markets that France still had a functioning government.

The unprecedented situation unfolded as Macron, who said he would wait until parliament was “structured” before making any decisions on a new government, was scheduled to leave the country on Wednesday for a Nato summit in Washington.

Attal had earlier said he would be willing to stay on in a caretaker role for as long as necessary to help oversee a smooth transition to a new government, if one could be found in a parliament split into opposing blocs. Macron asked him to remain “for the time being, to ensure the country’s stability”, the Élysée palace said.

Gaël Sliman, of the pollster Odoxa, asked: “Is this the biggest crisis of the Fifth Republic? Emmanuel Macron wanted clarification with the dissolution, now we are in total uncertainty. A very thick fog.” France’s Fifth Republic was established in 1958.

The Green leader, Marine Tondelier, one of a number of NFP figures seen as potential future prime ministers, said: “According to the logic of our institutions, Emmanuel Macron should today officially invite the NFP to nominate a prime minister.”

While the NFP’s leaders met again on Monday to try to agree a way forward, the leftist alliance appeared divided on how to proceed, with Jean-Luc Mélenchon, the leader of the radical-left France Unbowed (LFI), ruling out any coalition deal with centrists.

Mélenchon’s lieutenant Manuel Bompard confirmed the party’s line on Monday. “The president must appoint as prime minister someone from the NFP, to implement the NFP’s programme, its whole programme and nothing but its programme,” he said.

Raphaël Glucksmann, a moderate who led the Socialist party list in the European elections, said on Sunday that the NFP must be open to dialogue and compromise with other parliamentary groups to govern, but Bompard refused to engage with that possibility.

Tondelier said the prime minister could be someone from any of the NFP’s four member parties or an outsider. Olivier Faure, the Socialist party leader, said a name would be presented this week but declined to speculate about coalitions.

Very little of the NFP’s radical economic programme, which includes raising the minimum wage, reversing Macron’s pension changes and capping the prices of key goods, would win parliamentary approval without coalition support.

Prominent centrists including the former prime minister Édouard Philippe and the long-term Macron ally François Bayrou have said they would be in favour of a coalition agreement stretching from the moderate left to the centre right, but excluding Mélenchon’s LFI.

“We can no longer have one bloc against another – it can’t work like that any more,” Bayrou said on Monday. “French voters have told us that we have to abandon, as far as we can, government ‘against the rest’ for government ‘with the rest’.”

Yaël Braun-Pivet, an MP from Macron’s camp and the outgoing speaker of the lower house, said voters were telling her: “No one has an absolute majority, so you have to work together to find solutions to our problems.”

The left’s surprise victory came after an anti-far-right “republican front” formed to avoid splitting the vote in three-way races in hundreds of constituencies. Le Pen denounced the strategy as unfair, but a senior RN member said the party had work to do.

“We cannot carry on like this,” Bruno Bilde, an RN deputy from northern France, told Le Monde, arguing that the party could “complain all it likes about the unfair system but when so many candidates lose you have to question the candidates’ credibility”.

Jordan Bardella, the RN party president who had led the campaign, said he accepted his part in the “defeat” of the parliamentary elections, as well as the success of the European elections in which his party topped the poll in France.

For more than 50 years, whenever France held a parliamentary election, voters would know the next morning which party would be in government and with what political agenda.

This time it is different. After Emmanuel Macron called a surprise snap election, and after the shortest campaign in modern history, French people delivered a spectacular rush of tactical voting to hold back a surge of far-right support. The resulting political landscape is divided and the outcome is complicated. Macron will take time to let the dust settle, his entourage has said.

An alliance of parties on the left, the New Popular Front, surprised pollsters by coming first with a strong result of 182 seats. But it fell significantly short of the absolute majority of 289 that would allow it to instantly form a government. This means the eurozone’s second largest economy, which is also the EU’s biggest military power, is entering a period of uncertainty with no clear roadmap, less than three weeks before it hosts the Olympic Games.

It could take weeks of dialogue and potential coalition-building to come up with a government and a prime minister. But France – with a powerful president and conflictual political system where parties reach vicious standoffs – does not have a recent tradition of building coalitions.

The French parliament is now roughly split into three blocs. In the lead is the New Popular Front, which blindsided Macron and the opposition when it managed to swiftly and efficiently unite four weeks ago to counter the far right. It is a rainbow grouping that in parliament will run from the firmly leftwing France Unbowed (LFI), which has the greatest number of seats at 74, to the Greens, who increased their seats to 28, through to the more centrist Socialist party, which significantly increased its seats to 59.

The broad left alliance’s deliberately strident policy manifesto included capping prices of essential goods such as fuel and food, raising the minimum wage, reversing Macron’s increase in the pension age to 64 and imposing a wealth tax.

It took care not to push forward one leader during the campaign. Jean-Luc Mélenchon, the veteran leftwinger and firebrand orator who founded LFI, regularly made TV appearances during the campaign. But each time he did, other parties in the alliance would carefully state that he was not in charge and was not necessarily their choice for prime minister.

The Green leader, Marine Tondelier, on Monday called for a calming consensus figure to be proposed as prime minister. It is uncertain how the parties on the left will choose a figurehead, and who it could be.

Macron’s centrist grouping finished in second place on 168 seats, only 14 seats behind the New Popular Front. The centrists, previously in government, lost 80 seats amid voters’ clear anger and rejection of Macron. But the president’s entourage immediately pointed out that although they were reduced in number, they were still standing.

Beaten back into third place came Marine Le Pen’s far-right, anti-immigration National Rally (RN) and its allies with an unprecedented 143 seats.

The key takeaway of the election’s final round is the renewed strength of the French tradition known as the “republican front”, in which voters from all backgrounds group together in tactical voting to hold back the far right.

In the space of one week, the far-right party and its allies went from a first round in which it topped the vote in more than half of the constituencies in France and was within reach of forming a government, to being knocked back into third position in the second round.

The left is now fearful of writing off the far right too quickly. The RN improved from the 89 seats it won in June 2022 to a historic 126. It expanded its presence to new areas such as the Dordogne in the south-west. Le Pen said victory had simply been deferred. She will now focus on the presidential campaign of 2027. The party leadership will face a reckoning over its strategy and haphazard vetting of candidates, one of whom dropped out after photos circulated of her wearing a Nazi cap.

The Socialist party leader, Olivier Faure, recognised the fractured and bruised nature of French society after a divisive campaign. For many on the left, the immediate priority is to address key issues for Le Pen’s millions of voters, including the cost of living and poor access to public services in rural and peripheral areas. This is essential if the left is to present a credible alternative and continue to hold the back the far right’s slow but steady rise.

Constitutionally, it is possible for a group in parliament to govern without an absolute majority. But to do this, that group would have to ensure that opposition forces do not club together to form a majority of 289 to vote them down.

The first session of the new parliament is on 18 July. It may be that only then will possibilities for coalitions become fully clear.

The New Popular Front (NFP), a four-party left-green alliance, was the shock winner of Sunday’s French parliamentary election, returning 182 deputies to a 577-seat assembly now split between three large opposing blocs, none with a majority.

Here is a look at which parties make up the NFP, what it proposes, who its key figures are – and whether they may be able to continue to work together.

Donald Trump’s “preposterous” efforts to disavow Project 2025, a rightwing blueprint for a radical takeover of the US government if the former president is re-elected in November, have been derided by former Republican figures.

The Project 2025 plan includes calls for replacing civil servants with Trump loyalists, eliminating the education department, putting the justice department under the president’s thumb and banning the abortion pill.

Democrats have made concerted efforts to say the 900-plus page document from the conservative Heritage Foundation thinktank would be representative of a second Trump presidency.

But although it was written by former members of Trump’s first administration, and he regularly echoes its policies in his speeches, last week Trump tried to disown the initiative.

Posting on his Truth Social website, the presumptive Republican nominee claimed to “know nothing about Project 2025” and have “no idea who is behind it”.

He added: “I disagree with some of the things they’re saying and some of the things they’re saying are absolutely ridiculous and abysmal. Anything they do, I wish them luck, but I have nothing to do with them.”

Olivia Troye, a former White House adviser to Mike Pence who sat in on policy sessions during Trump’s first presidency, said Trump’s attempt to distance himself from Project 2025 was driven by a recognition that its deeply controversial policy prescriptions could sink his election bid.

“This is preposterous if you look at the collaborators and the authors of this plan,” she told CNN when asked whether Trump’s denial was credible. “A lot of these people…served in Trump’s cabinet during his administration. There are people that I worked with. I sat in those policy meetings with them.”

Troye identified various figures – including John McEntee, who was Trump’s director of White House personnel, Stephen Miller, a senior adviser in his first administration, Ben Carson, the housing and urban development secretary in his cabinet, and Ken Cuccinelli, a former deputy secretary of homeland security – as among the project’s leading architects.

Carson has been “out there on the campaign trail” with Trump, she noted.

“I think what this is telling us is that Donald Trump knows that what is written in this plan is so extreme that it is damaging to his possibility of getting elected, and that’s what he’s concerned about.”

“Exactly how do you ‘disagree’ with something you ‘know nothing about’ or ‘have no idea’ who is behind, saying or doing the thing you disagree with?” said former RNC Chairman and current MSNBC host Michael Steele in echoing Troye’s derision.

“And how exactly don’t you know that Project 2025 director Paul Dans served as your chief of staff at the Office of Personnel Management, and associate director Spencer Chretien served as your special assistant and associate director of presidential personnel?”

Among the plan’s more drastic proposals are to fire thousands of permanent civil servants and replace them with hired conservative Trump loyalists, dismantling the federal education department, asserting presidential power over the Department of Justice – which is nominally independent – and a ban on the abortion pill.

The Democrats, currently in the throes of a fierce internal debate over whether to retain Joe Biden as their presidential candidate, have settled on trying to make Project 2025 a household phrase in a drive to illustrate what a second Trump presidency would mean.

Troye said the project should be seen as a threat not just to Democrats but to moderate conservatives, too.

“If you go through and really read through this plan, this is complete overreach by the federal government on our individual liberties,” she said.

“[It talks] about law enforcement and how they’re going to use federal law enforcement in local states and local cities … with no oversight. Because there’s no oversight when they do that. They’ve learned all the lessons during the first Trump term, and that is what is frightening here. I think we need to be paying attention to this, and no amount of distancing by Donald Trump should be believed …I sat in [on] policy-making meetings with these people.”

Trump surrogates have tried to back up his effort to separate himself from the project, with Marco Rubio, a Republican senator from Florida who has been touted as a possible running mate for Trump, claiming in a Sunday interview with CNN that there was no connection between Project 2025 and Trump.

“Thinktanks do thinktank stuff. They come up with ideas, they say things,” he said. “But our party’s candidate for president is Donald Trump.”

He also dismissed the importance of comments by the Heritage Foundation’s president, Kevin Roberts, who said in an interview with Steve Bannon’s War Room podcast last week that conservative-driven “second American revolution” will be bloodless “if the left allows it to be”, viewed by many Democrats as an implied threat of political violence.

“He’s not running for president,” Rubio said. “Our candidate’s Donald Trump. I didn’t see Donald Trump say that.”

The denials appear to be undermined by close studies of the personnel involved in the document’s formulation.

Of the 38 people involved in the writing and editing of Project 2025, 31 of them were nominated to positions in Trump’s administration or transition team – meaning 81% of the document’s creators held formal roles in Trump’s presidency.

A Russian military court has sentenced a playwright and a theatre director to six years in prison on charges of “justifying terrorism” in a play about women marrying jihadists in Syria.

The judge sentenced director Yevgeniya Berkovich and writer Svetlana Petriychuk after moving their trial behind closed doors.

The women’s arrest in May last year sent shock waves through Russia’s artistic community, which has faced unprecedented pressure from the Kremlin since Russia sent troops to Ukraine.

The length of the sentence announced by the judge on Monday was the same as requested by prosecutors last week and one year less than the maximum possible.

Berkovich, 39, has written poems criticising Russia’s military offensive in Ukraine and her supporters said they believed the court case could be linked to this.

Rachel Denber, deputy director for Europe and Central Asia at Human Rights Watch, wrote on X that the women were sentenced “on utterly absurd charges, in an unfair trial that is blatant retaliation against Berkovich for speaking out against Russia’s war on Ukraine”.

Human rights group Amnesty International said the pair were “being targeted simply for exercising the right to freedom of expression” and called for their immediate release.

The two women were brought in for the sentencing wearing handcuffs and sat in a glass-walled dock in a courtroom heavily guarded by police with masked faces, AFP journalists saw.

Berkovich, wearing a white shirt, tried to smile and show a victory sign to supporters, while Petriychuk, 44, in a blue dress, looked tense.

Defence lawyer Ksenia Karpinskaya said after the sentence that “today was an absolutely illegal, unfair hearing” and that the women were “absolutely innocent”, to applause from those present.

“Of course we will appeal against this decision,” Karpinskaya said, condemning the “cruel sentence”.

“Today at the court hearing, Svetlana said that she … will never plead guilty for something she did not do. Yevgeniya said the same,” the lawyer said.

When the judge read out the women’s sentence, Berkovich’s husband, Nikolai Matveyev, who works for an independent theatre company, burst into tears.

Prosecutors charged the pair over their 2021 play Finist The Brave Falcon, about Russian women who were lured to marry Islamic State group militants in Syria and imprisoned upon returning to Russia. It was awarded two prestigious Golden Mask theatre awards.

The Kremlin has brought artistic institutions under tighter control since launching its Ukraine offensive in 2022. Many of Russia’s prominent artistic figures have left the country.

“I staged the play to prevent terrorism,” Berkovich said during the trial, denying the charges.

Among those who came to the court on Monday to support the women was Dmitry Muratov, a Nobel prize-winning newspaper editor, who had called for their charges to be dropped.

The women and their defence lawyers gave their final arguments in court in Monday during a closed hearing.

The judge ruled last month that the trial would continue behind closed doors after the prosecution said witnesses were being threatened on social media.

Media and supporters were only allowed to attend the sentencing.

A Russian military court has sentenced a playwright and a theatre director to six years in prison on charges of “justifying terrorism” in a play about women marrying jihadists in Syria.

The judge sentenced director Yevgeniya Berkovich and writer Svetlana Petriychuk after moving their trial behind closed doors.

The women’s arrest in May last year sent shock waves through Russia’s artistic community, which has faced unprecedented pressure from the Kremlin since Russia sent troops to Ukraine.

The length of the sentence announced by the judge on Monday was the same as requested by prosecutors last week and one year less than the maximum possible.

Berkovich, 39, has written poems criticising Russia’s military offensive in Ukraine and her supporters said they believed the court case could be linked to this.

Rachel Denber, deputy director for Europe and Central Asia at Human Rights Watch, wrote on X that the women were sentenced “on utterly absurd charges, in an unfair trial that is blatant retaliation against Berkovich for speaking out against Russia’s war on Ukraine”.

Human rights group Amnesty International said the pair were “being targeted simply for exercising the right to freedom of expression” and called for their immediate release.

The two women were brought in for the sentencing wearing handcuffs and sat in a glass-walled dock in a courtroom heavily guarded by police with masked faces, AFP journalists saw.

Berkovich, wearing a white shirt, tried to smile and show a victory sign to supporters, while Petriychuk, 44, in a blue dress, looked tense.

Defence lawyer Ksenia Karpinskaya said after the sentence that “today was an absolutely illegal, unfair hearing” and that the women were “absolutely innocent”, to applause from those present.

“Of course we will appeal against this decision,” Karpinskaya said, condemning the “cruel sentence”.

“Today at the court hearing, Svetlana said that she … will never plead guilty for something she did not do. Yevgeniya said the same,” the lawyer said.

When the judge read out the women’s sentence, Berkovich’s husband, Nikolai Matveyev, who works for an independent theatre company, burst into tears.

Prosecutors charged the pair over their 2021 play Finist The Brave Falcon, about Russian women who were lured to marry Islamic State group militants in Syria and imprisoned upon returning to Russia. It was awarded two prestigious Golden Mask theatre awards.

The Kremlin has brought artistic institutions under tighter control since launching its Ukraine offensive in 2022. Many of Russia’s prominent artistic figures have left the country.

“I staged the play to prevent terrorism,” Berkovich said during the trial, denying the charges.

Among those who came to the court on Monday to support the women was Dmitry Muratov, a Nobel prize-winning newspaper editor, who had called for their charges to be dropped.

The women and their defence lawyers gave their final arguments in court in Monday during a closed hearing.

The judge ruled last month that the trial would continue behind closed doors after the prosecution said witnesses were being threatened on social media.

Media and supporters were only allowed to attend the sentencing.

Labour is unlikely to rush into proscribing Iran’s Islamic Revolutionary Guards Corps, and will instead examine whether a new category of state-backed terrorism needs to be devised.

David Lammy, the foreign secretary, will also consult colleagues on the implications for Iranian foreign policy of the election at the weekend of a reformist-backed president, Masoud Pezeshkian.

An issue for the west is testing – either through private talks in Oman or more public dialogue – whether Pezeshkian has any real influence on foreign policy and wants to take steps to dial down Iran’s nuclear programme, which would make it possible for sanctions to be eased.

In opposition, Labour said it would proscribe the IRGC, a step that has caused deep concern in Tehran, with foreign ministry advisers warning that such a decision could have an untold damaging impact on UK-Iran relations.

The IRGC is an arm of the Iranian state, and it would be a major precedent for the UK to proscribe part of another state on the basis that it is involved in terrorist acts.

Speaking to the Guardian at the weekend, Lammy said: “We recognise there are real challenges from state-sponsored terrorist activity, and I want to look closely at those issues, and how the predecessor system works for states, as well as for specific terrorist organisations.”

His aides say he is examining a possible amendment to existing laws to allow the government to put targeted proscription-style restrictions on the operations of state-linked organisations such as the IRGC. But this might take time to develop, and the Labour manifesto referred to a possible new regime for state-sponsored terrorism without identifying the IRGC.

David Cameron, Lammy’s predecessor as foreign secretary, withstood wide parliamentary pressure to proscribe the IRGC, arguing such a step could mean Tehran cutting off diplomatic relations with the UK, something Lord Cameron was reluctant to do since he valued the direct, if often angry, conversations with the then Iranian foreign minister.

On the election of Pezeshkian, Lammy said: “I’ve seen the development over the last 24 hours. I want to speak to other colleagues about that and what that really means on the ground, I think it’s a little bit early to say.”

Many Iranian experts disagree about the extent to which the president can influence the direction of foreign policy, or whether ultimately major strategic decisions are reserved by those around the supreme leader, Ayatollah Ali Khamenei.

In a sign that Iran’s overall outlook will initially change only at the margins, Pezeskhian sent a letter to Hassan Nasrallah, the secretary general of the Iran-backed Hezbollah group, saying he was confident that the resistance movements in the region would “not allow [Israel] to continue its warmongering and criminal policies against the oppressed people of Palestine and other nations of the region”.

He also told the Russian president, Vladimir Putin, that Iran was ready to sign a delayed comprehensive cooperation agreement with Moscow.

There is speculation Abbas Araghchi will be the foreign minister in the new government. He worked as the deputy to the former foreign minister Javad Zarif, who acted as Pezeshkian’s campaign aide.

On Monday Lammy met the Canadian foreign minister, Mélanie Joly, in London, the first foreign minister to meet him in the capital since his appointment. Last month Canada announced it would proscribe the IRGC. It had also proscribed the IRGC’s overseas arm, the al-Quds force.

Ottawa already has no diplomatic relations with Iran, so the sacrifice is not comparable to the step the UK would take.

The effect of proscription is that members and supporters, both moral and financial, are criminally liable if they help a proscribed organisation. In a paper last year Jonathan Hall, the UK’s adviser on terrorism law, argued the effect of proscribing the IRGC would be to accept, contrary to the UK’s longstanding policy position, that state forces and therefore states can be “concerned in terrorism” within the Terrorism Act 2000.

He suggested the 2000 law might then need amending to make it legitimate for states to use force in line with humanitarian law.

A New Mexico judge decided on Monday that the actor Alec Baldwin’s role as co-producer is not relevant to the involuntary manslaughter trial over a fatal shooting on the set of the western film Rust.

The judge Mary Marlowe Sommer ruled that evidence would not be allowed at trial about Baldwin’s secondary role on the movie, siding with defense attorneys.

“I’m having real difficulty with the state’s position that they want to show that as a producer he didn’t follow guidelines and therefore as an actor Mr Baldwin did all of these things wrong that resulted in the death of Ms Hutchins because as a producer he allowed these things to happen,” Marlowe Sommer said. “I’m denying evidence of his status as a producer.”

The special prosecutor Erlinda Johnson argued unsuccessfully to allow evidence that Baldwin’s “role as a producer made him keenly aware of his responsibilities on set” for safety.

“It goes to Mr Baldwin’s knowledge, knowing that his conduct on set was negligent,” she said.

Baldwin sat between his lead attorneys Luke Nikas and Alex Spiro, with a yellow legal pad on the table in front of him.

The trial starts on Tuesday with jury selection and is scheduled to last 10 days.

Lawyers for Baldwin pushed for the case to be dismissed last month, arguing that FBI testing of the firearm had damaged the weapon before lawyers were able to examine it for possible modifications. The defense team alleged the gun was damaged at the time of the incident and accused prosecutors of withholding potentially “exculpatory evidence”.

Sommer rejected the dismissal request, saying Baldwin’s lawyers had not proved prosecutors acted in bad faith. But the judge also said prosecutors would have to disclose to the jury the “destructive nature of the firearm testing, the resulting loss and its relevance and import”.

Defense attorneys have asked the judge to exclude consideration of Baldwin’s secondary role as a co-producer on Rust, arguing it’s irrelevant to allegations of negligence, and might confuse jurors. Prosecutors disagree and say it was likely Baldwin’s imposing role as a producer that emboldened him to act recklessly and disregard the safety of others in allegedly flouting gun-safety protocols.

The defense team and prosecutors disagree about Baldwin’s contractual authority as producer over crew members who dealt with weapons and safety.

Prosecutors argue that a state workplace safety investigation, which found serious violations on set, was incomplete, untrustworthy and should be prohibited from the trial.

Baldwin is charged with a single felony count of involuntary manslaughter punishable by up to 18 months in prison if he is convicted.

Hannah Gutierrez-Reed, the armorer on set, was convicted of involuntary manslaughter in cinematographer Halyna Hutchins’s death and sentenced to 18 months in prison. She is appealing the conviction.

In October 2021, Baldwin was rehearsing a cross-draw maneuver with the revolver when the gun went off, killing Hutchins and wounding director Joel Souza.

Baldwin has pleaded not guilty and claims the gun fired accidentally after he followed instructions to point it toward Hutchins, who was behind the camera. Unaware the gun contained a live round, Baldwin said he pulled back the hammer – not the trigger – and it fired.

Baldwin’s attorneys also want to bar discussion at trial of the actor Brandon Lee’s death from a fatal shot to the abdomen while filming a scene from The Crow in 1993. In that instance, a makeshift bullet was mistakenly left in a gun from a previous scene and struck Lee while filming a scene that called for using blank rounds.

Prosecutors have agreed not to illicit testimony about The Crow, but also contend that Baldwin knew about safety risks posed by guns – even when live rounds are not present. Attorneys for Baldwin argue that it was inconceivable that live rounds would wind up on set.

Prosecutors want to exclude a letter signed by crew members that disputes the characterizations of the Rust set as chaotic or dangerous prior to the fatal shooting.

Prosecutors also want to exclude from trial the conclusions of the safety investigation into the fatal shooting that places much of the blame on assistant director Dave Halls. Halls has pleaded no contest to negligent use of a firearm and may be called to testify at Baldwin’s trial.

Rust Movie Productions paid a $100,000 fine to resolve violations of state safety regulations that were characterized as “serious” but not willful, under a 2023 settlement agreement. Prosecutors say conclusions of the investigation are easily contradicted by more reliable information.

Baldwin’s attorneys say the report cannot be ruled out as evidence and that state occupational safety officer Lorenzo Montoya should be allowed to testify at trial.

Another pretrial motion might defuse snipping between the prosecution and defense teams. Prosecutors want the judge to preclude accusations of “prosecutorial misconduct” and “personal attacks”.

Prosecutors also want the judge to exclude evidence and arguments designed to garner sympathy for Baldwin, including indications of remorse or the impact of events on his family, arguing that it has no bearing on determining guilt.

Andrew Tate and his brother, Tristan, have been accused of failing to pay tax on £21m of revenue from their online businesses.

Devon and Cornwall police are bringing a civil claim against the brothers and a third person, a woman referred to as J, over unpaid tax, Westminster magistrates court heard on Monday. The force is seeking about £2.8m in seven frozen bank accounts.

Sarah Clarke KC for Devon and Cornwall police said: “Andrew Tate and Tristan Tate are serial tax and VAT evaders. They, in particular Andrew Tate, are brazen about it.”

It is claimed that they paid no tax in any country on £21m revenue from businesses online earned between 2014 and 2022. Clarke quoted from a video posted online by Andrew Tate, in which the self-proclaimed misogynistic social media influencer said: “When I lived in England I refused to pay tax.”

The court heard he said his approach was “ignore, ignore, ignore because in the end they go away”. The court also heard that the brothers had a “huge number of bank accounts” in the UK, seven of which have been frozen.

Clarke said that money “washed around UK bank accounts”, that were used as a “mechanism for moving revenues from their business activities through a wide number of accounts”.

“That’s what tax evasion looks like, that’s what money laundering looks like,” she told the court.

The money came from products they sold online as well as their OnlyFans sites, the court was told.

The Tates are accused of failing to pay tax and of money laundering in both the UK and Romania, the court heard. It is claimed the brothers paid just under $12m (£9.3m) into an account in J’s name – and opened a second account in her name, even though she had no role in their businesses.

Devon and Cornwall police allege this was fraud by false misrepresentation.

Money from Cobra Tate, Hustlers’ University and War Room was paid into the first account, held with Stripe, and the majority of payments out went to one of Andrew Tate’s accounts, the court heard.

It was opened in February 2019 in J’s name with an incorrect date of birth, and later driving licences belonging to both Andrew Tate and J were submitted as proof of identity and address to the payment service provider – Tate in October 2019 and J in June 2022.

The proceedings are civil, which uses a lower standard of proof than criminal cases. Paul Goldspring, the chief magistrate, will decide on the balance of probabilities whether what the police claim is true.

J also moved money through her own account, including one payment of £805,000 that was made to her Revolut account, the court heard. Of this, £495,000 was paid to Andrew Tate, and £75,000 to an account in J’s name that was later converted to cryptocurrency, it is alleged.

Gary Pons, for J, argued that the funds in the Gemini account were in cryptocurrency and therefore could not be frozen at that time.

The case was adjourned until Tuesday.

People in Gaza City have reported one of the heaviest attacks by Israeli forces since 7 October, sending thousands of Palestinians fleeing from an area already ravaged in the early weeks of the nine-month-old war.

The latest Israeli incursion into the eastern sector of Gaza City came as Israel’s far-right coalition parties threatened again to stop ongoing negotiations in Qatar for a ceasefire, arguing that halting the fighting now would be a huge mistake.

Sporadic militant activity has hindered Israeli efforts to maintain control in Gaza. Despite claiming authority over the Gaza City area months ago, Israel has had to contend with persistent pockets of resistance, forcing a re-evaluation of its military strategy and redeployment of forces.

Before the recent offensive, the Israeli military said it had issued evacuation orders in the targeted zone.

The territory’s civil emergency service told Reuters it believed that dozens of people had been killed in Gaza City but that teams were unable to reach them because of offensives in a number of areas.

Local sources said Israeli warplanes had bombed a residential apartment near an industrial junction south of Gaza City, killing two citizens and wounding five others. They said a house south of Gaza City was bombed, killing one person and wounding seven others.

“Eyewitnesses said that thousands of citizens were displaced from areas south-west of the city towards the north-west and spent the night on the streets without shelter,” a source said.

Sayeda Abdel-Baki, who was sheltering at her relatives’ home in the Daraj neighbourhood of Gaza City, told Associated Press: “We fled in the darkness amid heavy strikes. This is my fifth displacement.”

Medics at al-Ahli Arab Baptist hospital in Gaza City had to evacuate patients to the already crowded and under-equipped Indonesian hospital in the northern Gaza Strip, Palestinian health officials said.

According to an Israeli military statement, Israeli forces were operating “following intelligence indicating the presence of Hamas and Islamic Jihad terrorist infrastructure, operatives, weapons and investigation and detention rooms, including in the Unrwa headquarters”.

Philippe Lazzarini, the commissioner general for Unrwa, the UN’s agency for Palestinian refugees, has previously criticised Hamas and Israel for occupying and using its facilities during the conflict.

Hopes among Gaza residents of a pause in the fighting had revived after Hamas accepted a key part of a US ceasefire proposal, prompting an official in the Israeli negotiating team to say there was a real chance of a deal.

However, the Israeli finance minister, Bezalel Smotrich, who heads a pro-settler party that is part of Benjamin Netanyahu’s governing coalition, said stopping the war now would be a huge mistake.

“Hamas is collapsing and begging for a ceasefire,” Smotrich wrote on X. “This is the time to squeeze the neck until we crush and break the enemy. To stop now, just before the end, and let him recover and fight us again, is a senseless folly.”

Concerns are growing in Israel over the substantial influence wielded by the far-right opponents of a ceasefire deal in the coalition.

Netanyahu is facing criticism from opposition parties, media and families of Israeli hostages, who accuse him of undermining efforts to reach a ceasefire and secure the release of the hostages for his own political survival.

On Monday, a senior Hamas official accused Netanyahu of stepping up combat and bombardment in Gaza in order to derail the latest truce effort. “Whenever a round of negotiations begins and a breakthrough is within reach, he disrupts it all and escalates the aggression and massacres against civilians,” the Hamas official told Agence France-Presse.

Netanyahu’s popularity plummeted after the 7 October attack by Hamas, which exposed serious flaws in Israeli security. Most political observers say he would lose elections if they were held now.

This is not the first time that far-right parties within the coalition have interfered with negotiations.

Dahlia Scheindlin, a political analyst and author of The Crooked Timber of Democracy in Israel, said: “It’s natural that the far-right parties have influence over Netanyahu’s decisions. They are his coalition partners and they were voted into that position. It would be strange if they didn’t have influence over Netanyahu. In addition, the war cabinet is gone and Netanyahu has a poor relationship with his defence minister and is suspicious of the military establishment. Any influence they have is certainly matched by the partners Netanyahu chose for his government.

“The unusual background is that Netanyahu didn’t really have a choice of coalition partners in 2022 because he had been indicted … So no other parties would join his government. When people elected this government in 2022, they weren’t expecting a war of this magnitude … they didn’t know that they were choosing leaders for this purpose.”

On Sunday, Netanyahu’s office issued a document saying any deal must allow Israel to resume its offensive “until it reaches its war goals”.

The document was heavily criticised by the opposition leader, Yair Lapid, who said: “What good does this do? We are at a crucial moment in the negotiations, the lives of the hostages depends on this. Why make such taunting announcements? How does this help the process?”

AFP, AP and Reuters contributed to this report

Scientists have for the first time recovered DNA from the remains of dingoes between 400 and 2,700 years old to find the predator’s population was well established across the continent thousands of years ago.

According to the researchers, modern dingoes share little genetic ancestry with domestic dogs introduced into Australia from Europe, but are instead descended from ancient dogs and wolves from China and the Tibetan plateau. Dingoes were closely related to modern New Guinea singing dogs, the research confirmed, with both sharing a common ancestor.

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Dingoes arrived in Australia between 3,000 and 8,000 years ago, the research published in the journal Proceedings of the National Academy of Sciences said, and probably came with Pacific traders on boats.

Scientists gathered ancient DNA from the remains of dingoes held in museums, recovered from Indigenous sites around Sydney and from caves in South Australia and south-west Western Australia.

Dr Sally Wasef, an expert on ancient DNA from the Queensland University of Technology and a lead author of the research, had the job of cutting the ancient specimens – mostly bone and teeth – for DNA analysis.

She said she was amazed when the results of carbon dating found several of the east coast specimens were between 700 and 2,700 years old.

“I thought they were fresh samples, so I was really shocked,” said Wasef, who has done previous genetic work on Indigenous remains, Egyptian mummies and world war victims.

“We have to respect these ancient remains because they are telling us a story,” she said.

The DNA analysis, including 42 ancient specimens, found the dingo established two distinct regional populations, split roughly along the Great Dividing Range.

Wasef said previously, it had been thought this division had formed during post-colonial human activity.

Dr Yassine Souilmi, of the University of Adelaide’s Australian Centre for Ancient DNA and Environment Institute and also a lead author of the study, said two genetically distinct groups also pre-dated the rabbit and dingo proof fence.

“We know that distinction goes back at least 2,500 years,” he said.

“We hope people making decisions about dingoes today now see these amazing animals have been around a long time, and had time to harmonise with the environment.”

The modern K’gari dingoes analysed had no domestic dog ancestry.

Genetic analysis found the ancient dingoes found in an Indigenous rock shelter at Curracurrang and modern dingoes living in alpine areas of Victoria and southern NSW were especially close to the New Guinea singing dog.

The ancient dingo remains from Curracurrang and Sydney’s Balmoral Beach had been held at the Australian Museum, the former since the 1960s.

Prof Jane Balme, of the University of Western Australia and a co-author, said those ancient dingoes were likely companions to Aboriginal people and at Curracurrang had been deliberately buried in a rock shelter.

Prof Alan Cooper helped recover samples from caves in South Australia’s Nullarbor which, he said, were littered with mummified dingoes that had either fallen into pitfall entrances or pursued prey, such as kangaroos, into the caves.

Cooper, an evolutionary biologist at Curtin University, said: “There are thousands of caves and probably thousands of dingoes. Because it’s so dry they mummify, and are really well preserved as a result.

“You get beautifully laid out specimens and you can choose dense bones and get very good genetic information from them.”

He said the study raised an intriguing question of why there was so little mixing between dingoes in the west and in the east, but also suggested there were likely two waves of dingo introductions into Australia.

Cooper is trying to work out if the introduction of the dingo into Australia pushed the thylacine and Tasmanian devil to extinction on the Australian mainland – a question which the study can not rule out.

Dingoes are sometimes referred to as wild dogs in the context of controlling their numbers. Ongoing DNA studies have shown almost all dingoes retain their ancestry, rather than being hybrids.

Prof Mike Letnic, a dingo expert at the University of New South Wales but who was not involved in the study, said the findings were “great news” because they “put to bed the idea that dingoes are hybrids with no conservation value”.

“The results add weight to efforts to conserve dingoes because it shows that they are a distinct group and that there has been much less hybridisation than was previously thought.”