The executive, Jim Keller, is moving to chipmaking giant Intel, both companies said Thursday.
Keller's departure adds to the challenges of the electric car giant's effort to craft self-driving technology.
Keller's last day at Tesla was Wednesday, and he was the third Autopilot project head to leave in about two years. He oversaw low-voltage hardware, autopilot software, and "infotainment," according to Tesla.
"We appreciate his contributions to Tesla and wish him the best," the auto firm said in a statement.
Pete Bannon, whose background includes working on Apple computer chips, was named the new head of Autopilot hardware. Responsibility for Autopilot software was given to Tesla director of artificial intelligence Andrej Karpathy
"Tesla is deeply committed to developing the most advanced silicon in the world and we plan to dramatically increase our investment in that area while building on the world-class leadership team we have in place," the company said.
Keller next week will start his job as an Intel senior vice president leading silicon engineering, according to the chip maker.
The move comes with US safety investigators probing a March 23 accident in which a Tesla Model X operating on Autopilot collided with a highway barrier near the town of Mountain View in California, catching fire before it was struck by two other cars.
The driver, identified as a 38-year-old man, Wei Huang, later died in hospital.
Tesla said this month it "withdrew" from a party agreement governing a National Transportation Safety Board probe.
But the NTSB announced the "removal of Tesla," after the company disclosed information the agency said could taint the public understanding of what happened, in violation of the agreement.
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