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KYIV, Ukraine — President Volodymyr Zelensky dismissed the head of Ukraine’s security services and its prosecutor general on Sunday, later announcing that hundreds of criminal investigations for suspected “treason and collaboration activities” were underway in the besieged country.

Ivan Bakanov, head of Ukraine’s security services, the SBU, and Iryna Venediktova, the country’s prosecutor general, were relieved of their posts Sunday in presidential decrees that announced the government’s biggest shake-up since Russia’s full-scale invasion began nearly five months ago.

As of Sunday, Ukraine had registered 651 criminal proceedings against employees across several high-level offices for allegedly collaborating with Russians or working against the nation’s goals, Zelensky said in his nightly address shortly after announcing the dismissals.

“Such an array of crimes against the foundations of the national security of the state and the connections detected between the employees of the security forces of Ukraine and the special services of Russia pose very serious questions to the relevant leadership,” Zelensky said.

He added that any employees working against Ukraine would be held accountable, and that he had already replaced the security heads of the Kharkiv and Kherson regions.

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During his remarks, Zelensky said agencies cannot have full functionality without an “effective leader.” He did not disclose in his address whether Bakanov or Venediktova were being investigated for alleged treason or other crimes.

As Ukrainian officials investigate what Zelensky called “specific actions and any inaction,” the nation is on heightened alert for airstrikes from its invader. Russian forces have stepped up their attacks beyond the front lines. The Pentagon estimates that up to 150 Ukrainians have been killed in Russian attacks on civilian areas in the past two weeks.

Russian Defense Minister Sergei Shoigu on Saturday ordered his forces to intensify attacks “in all operational sectors” of Ukraine. As of Sunday night, Zelensky said, Russia had used more than 3,000 cruise missiles in Ukraine.

“It is impossible to count the number of artillery and other projectiles that were used against our country and our people,” he said. “But it is definitely possible to bring all Russian war criminals to justice.”

As Ukraine braced for more strikes, Sunday’s dismissals highlighted distrust within Zelensky’s ranks.

Bakanov’s leadership of Ukraine’s domestic intelligence and security agency had been under criticism since the start of the war after three former officials in the SBU were charged with state treason in late March.

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Two of the officials, Gen. Serhiy Kryvoruchko and Col. Ihor Sadokhin, had worked in the Kherson office. Kherson was the first major city captured by the Russians and was taken with little resistance — in large part because Ukrainian troops did not blow up the Antonovskiy Bridge, which connects the city to an area from which Moscow-backed forces had advanced.

Appointed to the top SBU role in 2019, Bakanov was a childhood friend of Zelensky’s. He ran his presidential campaign and, before that, his entertainment company. Ukrainian opposition parties criticized his appointment, saying he was not qualified to lead the SBU. Bakanov has a background in law and economics, according to his agency biography.

Zelensky had been looking to replace Bakanov and other security officials for weeks, Politico reported. He did not name a replacement for Bakanov in Sunday’s decree or his nightly address.

A former adviser to Zelensky and a lawmaker from his party, Venediktova was appointed as prosecutor general in 2020. Since shortly after Russia’s full-scale invasion began in February, she and her office have been investigating suspected war crimes.

Venediktova, the nation’s first female prosecutor general, has been profiled in several prominent U.S. news outlets for her prosecution of alleged Russian atrocities. She told Washington Post columnist David Ignatius in late May that her office had more than 13,000 war crime cases on Russian atrocities, including civilian killings, rape and torture.

She has faced criticism from some Ukrainians who have said she has not achieved sufficient results in high-profile anti-corruption cases.

Venediktova succeeded former prosecutor general Ruslan Ryaboshapka. When Ryaboshapka was fired, Zelensky had said he had failed to produce results.

In 2020, Zelensky said another person would be hired if Venediktova also could not yield results, according to the Kyiv Post.

Venediktova will be replaced by Oleksiy Symonenko, according to Zelensky’s decree announcing her dismissal.

As Ukraine continues to evaluate the actions of its officials, Zelensky said, the questions about its leadership will receive a “proper answer.”

Somasundaram reported from Washington.