Princeton turned the football over four times in its game against Penn, which completely erased an otherwise dominant offensive performance in which the Tigers rolled up 29 first downs and 444 total yards.
Penn quarterback Billy Ragone ran for one touchdown and also hit tight end Ryan O'Malley with a touchdown pass as the Quakers downed Princeton, 28-21.
Princeton again employed the two-quarterback system, with Quinn Epperly running the ball as many times as he completed passes, but Connor Michelsen's (28-of-46 for 272 yards) three interceptions gave the Quakers the opportunities they needed to score 14 unanswered points in the fourth quarter.
The Quakers relied heavily on their rushing attack, running the ball 47 times for 211 yards on the ground.
The game's three leading rushers were all Quakers, with running back Lyle Marsh (104 yards), quarterback Billy Ragone (61 yards) and running back Jeff Jack (50 yards) all outpacing any Princeton player.
Its running game ineffective, Princeton turned to the arm of Connor Michelsen as the Tigers utilized 10 different pass receivers. Wide receivers Matt Costello (seven receptions), Roman Wilson (five receptions) and Shane Wilkinson (five receptions) led the way.
With the loss, Princeton dropped to 4-4 on the season, but more importantly, it now has a 3-2 Ivy League record. The victory raised the Quakers to 4-4 overall, 4-1 in the Ivy League.
The Tigers must now look for help in order to take the Ivy Conference, but they will need to focus on their final two Ivy opponents. Princeton will travel to Yale on Nov. 10 and finish the season at home when it hosts Dartmouth on Nov. 17.