The 54-year-old was detained in Moscow in 2018 -- allegedly with a cache of classified documents -- and then sentenced to 16 years in a remote Russian penal colony for espionage.
Before Turkey's government announced his release as part of a major prisoner swap Thursday, Whelan was losing hope of ever being freed, having seen other high-profile Americans including basketball star Brittney Griner sent home through negotiations.
"The Russians have ruined my life. One that is destined to end in a slave labor camp, fraught with intolerable conditions," he told his parents in December 2023, after the Kremlin announced it rejected an offer to swap him.
Both his detention and release came against the backdrop of the worst crisis in ties between Washington and Moscow in decades, exacerbated by the Kremlin's large-scale military intervention in Ukraine.
Whelan's case was then overshadowed in March 2023, when the Kremlin arrested Evan Gershkovich, a US journalist working in Russia for The Wall Street Journal -- also on espionage charges.
Both were freed in Thursday's prisoner exchange.
- Mr Bean, not Mr Bond -
Whelan was born in Canada in 1970 to British parents of Irish origin and he holds British, American, Irish and Canadian citizenships.
He grew up in the US state of Michigan, where he studied criminal justice before becoming a police officer and joining the military.
He served two tours in Iraq in 2004 and 2006 but was discharged when the Marine Corps ruled he had tried to embezzle thousands of dollars of government funds.
Whelan was the security director for a US automotive parts manufacturer at the time of his arrest in Moscow, where he said he had travelled for a wedding of another US veteran.
He was detained by Russia's notorious FSB security service after he took a flash drive from an acquaintance, thinking it contained holiday photographs.
Moscow claimed he had illegally taken possession of state secrets that could harm national security.
"Russia thought they caught James Bond on a spy mission; in reality, they abducted Mr Bean on holiday," Whelan said in 2019.
That was one of a series of vocal protests Whelan made against his detention alongside animated appeals to US officials for his release. Throughout his trial and detention, he maintained his innocence.
At one hearing, most of which were held behind closed doors, Whelan called on then-US president Donald Trump to secure his release to "keep America great" -- a reference to a popular Trump slogan.
Washington had designated him as "wrongfully detained."
Whelan was first held in Moscow's infamous Lefortovo prison, which has housed Russia's most high-profile prisoners, including Gershkovich, who worked for AFP.
Whelan complained in detention that he was refused sufficient treatment for persistent health problems and that he had been injured by prison guards.
And when he was moved to an isolated penal colony in the region of Mordovia after being convicted, concerns grew over his conditions. He was tasked with sewing clothes in a prison factory in icy temperatures.
He claimed to have been assaulted by another inmate for being an American citizen and punished by prison staff for refusing an interview with state-run media RT, known as Russia Today.
- 'Rattled like never before' -
Speculation that he could be included in a prisoner swap between Moscow and Washington grew immediately after Whelan's conviction in June 2020.
His family said he decided not to appeal the verdict, hoping instead for a speedier exchange.
But he was not included in a swap two years later when another former US marine, Trevor Reed, was released in April 2022 after being convicted of drunkenly assaulting a police officer in Moscow.
And Whelan was also left out of an exchange in December 2022, when Griner -- arrested at a Moscow airport and sentenced to nine years on minor drug offenses -- was returned to the United States in exchange for arms dealer Viktor Bout.
A US envoy who spoke to Whelan after Griner's release said the conversation was "one of the toughest phone calls" he had ever made.
Griner vowed to lobby for Whelan's release, and Biden said he would never give up on him.
But when Gershkovich was detained in March 2023, Whelan grew increasingly concerned he would be looked over once more.
"Paul seems rattled like never before, understandably apprehensive that the US government will choose not to bring him home again, now that there is another American wrongfully detained," his brother said.
On Thursday, it appeared his fears could finally be laid to rest.
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