Billy Ragone passed for one touchdown and ran for another while the Pennsylvania defense forced four turnovers — including three interceptions — as the Quakers outdueled Princeton 28-21 Saturday in a battle of first-place Ivy League foes.

Penn (4-4, 4-1) is still tied with next Saturday's opponent, Harvard, for first atop the Ivy League.

Each of Ragone's scores catapulted the Quakers into the lead, including a game-winning, 3-yard TD run with 2:59 to play that snapped a 21-21 tie.

Ragone's other score, a 14-yard TD pass to Ryan O'Malley, put the Quakers ahead 14-13 in the second quarter.

The Penn defense intercepted Princeton's Connor Michelsen three times, with C.J. Mooney returning the final pick for a 15-yard score.

The turnovers undermined an effective offensive outing for the Tigers (4-4, 3-2), who out-passed Penn 319-96.

Penn's Lyle Marsh ran for 104 yards on 19 carries.