Up two with time winding down on its next drive, Harvard came up one yard short of what would have been its 34th first down on Juszczyk's 15th reception—second-best in team history—and was forced to give the ball back to the Tigers.

It was on Princeton's final drive on which the most costly Harvard miscues came.

The Tigers had a first down at their own 37 when backup quarterback Quinn Epperly—who had replaced an injured Michelsen—was sacked, ticking precious seconds off the clock. But senior John Lyon was called for an excessive celebration penalty, turning a 2nd-and-17 at the Princeton 30 into a 1st-and-10 at the Tigers 45.

Two plays later, Epperly made a poor read, throwing the ball directly at junior safety Chris Splinter. But the safety could not hold on to what would have been a game-ending interception for the second time in the contest, and after failing to put the game away again, the Crimson soon learned that Splinter’s drop would be the straw that finally broke the camel’s back.

Within seconds, Epperly had found Roman Wilson on a 39-yard fade to the back corner of the end zone, giving Princeton its fourth touchdown in 11 minutes and its 29th unanswered point to clinch the upset and leave the Crimson stunned.

The Tigers had won a match in which it had given up 634 yards of offense, thanks in large part to three Crimson turnovers and 12 Harvard penalties, compared to just five for Princeton.

Harvard, meanwhile, had lost a contest in which its quarterback broke the all-time Crimson passing record with 448 yards and tied the all-time record with five touchdowns, in which Scales eclipsed 100 yards on the ground, and in which Jusczcyk, Brate, and Berg all registered over 100 through the air.

The contest followed a similar script to last year’s game between the two teams, when Harvard went up 26 before a dramatic Tigers second-half rally cut the Crimson lead to 42-39. But last season, Harvard answered back with two touchdowns to put the game away and keep its undefeated league season rolling. This year, a much-improved Princeton squad was able to finish the comeback and take first place for itself.

“[Harvard is] probably the better team,” Princeton coach Bob Surace said. “But we got some breaks, and we’re very fortunate.... I’m glad we didn’t play them in a seven-game series, but it’s every given Sunday—or Saturday.”

—Staff writer Scott A. Sherman can be reached at ssherman13@college.harvard.edu.