Ahmed Khalfan Ghailani, from Tanzania, is accused of helping al Qaeda to carry out the 1998 bombings on U.S. embassies in Tanzania and Kenya which killed 224 people.
Ghailani was captured in Pakistan in 2004 and taken to Guantanamo in 2006, where his lawyers say he was tortured.
Other detainees have been tried in military commissions, but Ghailani is the first to be tried in the civilian courts.
If the trial is successful, it could pave the way for the Obama administration to prosecute Khalid Sheikh Mohammed, the alleged mastermind behind the 9/11 attacks.
It is also a test case for the U.S. President's pledge to close the military base in Cuba by next January.
Ghailani, said to be in his 30s, is accused of purchasing a truck and explosives used in the Tanzania attack and of conspiring with Osama bin Laden and other members of al Qaeda to kill Americans.
Jonathan Turley, law professor at George Washington University, said: 'Ghailani is one of the lucky ones who will be given a real trial in front of a real judge.
'The Obama administration seems to be suggesting that they are willing to give this a try and if successful, they might allow it for trial of other people.'
Ghailani was subject to what the government referred to as 'enhanced interrogation' at secret prisons run by the CIA before he was moved to Guantanamo.
His lawyers said he endured torture. He had pleaded not guilty to the charges at a hearing last year.
Yesterday, U.S. marshals escorted him from Cuba to New York for the trial.
Critics of trying terrorism suspects in civilian courts have said it could lead to the leaking of government secrets and any evidence obtained through torture could be inadmissible.
Mr Turley said: 'He's charged with a crime that occurred 12 years ago. That's a very long time to go without a trial.
'Witnesses become scarce, evidence become stale and memories fade.'
Ghailani faces life in prison if convicted.
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