Vladimir Putin has ordered the mobilisation of army reservists to support Moscow’s ailing campaign in Ukraine and warned that he would use Russia’s nuclear arsenal if its “territorial integrity” was “threatened”, declaring: “This is not a bluff.”

The warning, which sparked immediate alarm in Washington and elsewhere, came as Putin claimed the west wanted to “weaken, divide and destroy Russia” and pressed on with plans to annex swaths of Ukrainian territory.

US president Joe Biden hit out at Putin’s “overt nuclear threats against Europe [and] reckless disregard for the responsibilities of a non-proliferation regime”, attacking the referendum and mobilisation plans as “outrageous acts”.

He added, in a speech to the UN General Assembly: “This war is about extinguishing Ukraine’s right to exist as a state, plain and simple . . . That should make your blood run cold.”

Moscow announced the “partial mobilisation” of 300,000 reservists ahead of heavily stage-managed votes in four occupied regions of Ukraine to join Russia.

Western officials have estimated that at present there are between 150,000 and 190,000 Russian forces on the ground in Ukraine.

The results of the “referendums”, starting on Friday, in the Russian-controlled areas of the Donetsk, Luhansk, Kherson and Zaporizhzhia regions are in effect a foregone conclusion, with the Kremlin in charge of the vote. It insists more than 80 per cent of the population in Kherson and Zaporizhzhia and more than 90 per cent in Donetsk and Luhansk want to join Russia.

Putin said: “Russia can’t give up people close to her to be torn apart by executioners and fail to respond to their desire to determine their own fate.”

In his reference to Russia’s nuclear capability, he added: “If its territorial integrity is threatened Russia will use all the means at its disposal.”

Nato secretary-general Jens Stoltenberg attacked Putin’s “dangerous and reckless nuclear rhetoric”, adding: “He knows very well that a nuclear war should never be fought and cannot be won, and it will have unprecedented consequences for Russia.”

Other western officials said the rapid annexation plan and partial mobilisation were signs of Putin’s weakness.

Biden said the Russian invasion of Ukraine violated the United Nations charter, promising to continue supporting Kyiv’s efforts to defend itself.

Russia appeared to acknowledge that advanced western weaponry such as the US-manufactured Himars rocket launcher system had begun to turn the tide on the battlefield.

“We are not at war with Ukraine, but with the collective west,” defence minister Sergei Shoigu said in a state television interview.

He added Moscow would only call up reserves, rather than deploy the conscript army. This would add 300,000 people to Russia’s fighting force, he said, asserting that those called up would have combat experience and military specialisations.

“These are not people who’ve never seen or heard anything about the army,” he said. “We’re not talking about the mobilisation of any students . . . They can calmly keep going to class,” Shoigu said. He claimed that after the reserves were called up, Russia would still have only used up 1 per cent of its mobilisation potential.

Russia’s invasion has faltered badly, with invading forces taking heavy casualties before retreating from central Ukraine in the spring and then ceding huge swaths of territory in the eastern Kharkiv region following a Ukrainian counter-offensive this month.

Additional reporting by Roman Olearchyk in Kyiv and Felicia Schwartz in New York

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