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Chief of Staff Gen. T. Michael Moseley and Secretary Michael W. Wynne resigned over the department's concern over two incidents, including the August flight of a B-52 bomber that flew across the country with nuclear weapons.
"Focus of the Air Force leadership has drifted" in terms of handling nuclear weapons and equipment, Defense Secretary Robert Gates said.
Gates also cited this year's discovery that four nuclear warhead fuses were accidentally shipped to Taiwan in 2006.
Gates said he asked for the resignations after consulting President Bush.