Israeli agents acting at the behest of American officials assassinated al-Qaida's second-in-command in August, in a brazen drive-by shooting in Iran's capital, according to a senior U.S. official.

Abdullah Ahmed Abdullah, whose nom de guerre was Abu Muhammad al-Masri, was killed along with his daughter, Maryam, as they were driving in an upscale Tehran neighborhood, according to the official, who spoke on the condition of anonymity because of the matter's sensitivity.

The operation, first reported Friday by the New York Times, further slims the leadership ranks of al-Qaida and removes the accused mastermind of the 1998 bombings of two U.S. embassies in East Africa that killed more than 200 people.

Al-Qaida has not officially acknowledged Masri's death, and no country has claimed responsibility. The Israelis coordinated the operation with the CIA, according to the official. The CIA, FBI and Pentagon declined to comment, and the White House did not respond to a request for comment. The Israeli prime minister's office and intelligence ministry also declined to comment.

Iran's Foreign Ministry issued a statement Saturday denying the report and the presence of any Qaeda members in Iran, accusing the United States and Israel of trying "to draw a link between Iran and such groups through falsification and the leakage of fabricated information to the media."

A report about Masri's death was posted and quickly deleted from a private al-Qaida forum last month, according to SITE Intelligence Group, which tracks extremism online. al-Qaida has repeatedly failed to acknowledge the deaths of its leaders in the past two years, fearful that to do so would betray its weakness, SITE founder Rita Katz said.

Iran, a Shiite Muslim theocracy whose ideology is at odds with that of Sunni Islamic groups such as al-Qaida, has long denied it has harbored al-Qaida, though top leaders such as Masri fled from Afghanistan after the 2001 terrorist attacks and sought refuge there. The Washington Post first reported in 2003 that Masri was in Iran.

After the Aug. 7 attack on Masri, Iran concocted a cover story. Its semiofficial news agency Fars reported that a Lebanese man, Habib Daoud, and his daughter, Maryam, were shot and killed by a rider on a motorcycle, according to SITE. Other Iranian news outlets said the man was a history professor.

In fact Habib Daoud was Masri, 57, an Egyptian-born former professional soccer player who became one of Osama bin Laden's most trusted lieutenants.

Masri joined al-Qaida at its inception in 1988, having fought the Soviets in Afghanistan. He ran al-Qaida's terrorist training camps and played a role in the group's mass-casualty attacks against Americans, including helping train the Somali militants who attacked U.S. forces in Mogadishu in 1993.

He was still on the FBI's Most Wanted Terrorists list as of Saturday, and the State Department offered a reward of up to $10 million for information leading to his arrest.

"Abu Mohammed al-Masri was among the most important - and culpable - al-Qaida leaders on the planet," said Nicholas J. Lewin, a former federal prosecutor in Manhattan who helped lead the investigation and prosecution of several of the group's senior leaders and operatives. "His death deprives al-Qaida of one of its most experienced and respected leaders."

Masri's daughter, Maryam, married bin Laden's son, Hamza bin Laden, who was being groomed for a leadership role and had also lived in Iran. He was killed last year in a U.S. counterterrorism operation in the Afghanistan-Pakistan region.

While it might seem strange that Iran would harbor senior al-Qaida leaders, Iran in the past had aided Sunni militant groups such as Hamas and Egyptian Islamic Jihad, and Osama bin Laden once contemplated Iran and al-Qaida teaming up against the West, counterterrorism expert Ali Soufan wrote in an article for the Combating Terrorism Center at West Point.

Many of al-Qaida's senior commanders have been sheltered in Iran, though one by one, they have been killed in recent years. With Masri's death, the only remaining member of al-Qaida's shura council - its core leadership - with operational al-Qaida terrorist experience is Saif al-Adel, who is believed still to be in Iran.

Al-Qaida's leader, Ayman al-Zawahiri, who assumed the mantle after the 2011 killing of Osama bin Laden by U.S. Special Forces in Pakistan, is 69 and ailing.

"Al-Masri's death leaves limited choices for al-Qaida to select a new emir with the legitimacy of a bin Laden or Zawahiri,'' Soufan said in an email. Adel is a prime contender, he said, but noted that "could be controversial because his appointment could leave al-Qaida open to accusations that it is acting as a puppet of the Iranian regime, whether or not these claims have any basis in reality."

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