Princeton players celebrate as fans storm the field following the Tigers' shocking 39-34 upset over the Harvard football team on Saturday afternoon in Princeton, N.J.

Up 34-10 two minutes into the fourth quarter, the Harvard football team seemed to be on its way to a 15th straight win. 29 unanswered Princeton points later, the Crimson’s streak had come to an end as the Tigers faithful stormed the field at Princeton Stadium following the Tigers' improbable 39-34 comeback victory.

With under a minute to go, Princeton starting quarterback Connor Michelsen was knocked out of the game with the Tigers (4-2, 3-0 Ivy) down two points. But backup QB Quinn Epperly threw up a prayer to the back corner of the end zone on a third and two from the 36-yard line, and Roman Wilson came down with the ball to put the Tigers up five with 13 seconds to go. After accruing 634 total yards of offense in the game, the Crimson didn’t have anything left after that, suffering its first Ivy loss since 2010.

"The thought of losing this season never crossed my mind until they actually scored that final touchdown," senior tight end Kyle Juszczyk said. “It’s tough; it’s probably the toughest loss I’ve ever had in my career.”

Over the course of the game, Juszczyk amassed 192 yards to go with three scores while Harvard quarterback Colton Chapple passed for over 448 yards, breaking the Crimson all-time record.

Harvard (5-1, 2-1) had seemed in control of Saturday afternoon's game in the first half, outgaining the Tigers 415-51 in total yards, but the tables turned in the second half. Princeton put together drive after drive in the fourth quarter, often taking advantage of good field position provided by the defense and special teams to help diminish and eventually eliminate what was a 24-point lead.

"[They were] extraordinarily different halves.... The best way I can explain it is ‘Go figure,'" Harvard coach Tim Murphy said. "We just couldn’t get it done when we had to."

After being shut out in the first two quarters, Princeton came to life in the second, beginning with an 86-yard drive that culminated in a one-yard touchdown run to make the score 20-7.

Freshman Paul Stanton fumbled the ensuing kickoff, setting up the Tigers at the Harvard four. But the Crimson defense held strong as Princeton settled for a field goal.

Juszczyk then brought in his third touchdown catch of the day to cap a 75-yard response by Harvard that gave it a 27-10 lead. An interception by senior linebacker Alexander Norman on Princeton’s next drive led to a touchdown reception by junior Cam Brate that put the Crimson up three scores.

Princeton returned the ensuing kickoff into Harvard territory, and four plays later Michelson connected on a 29-yard touchdown pass. Epperly then completed a pass for a two-point conversion to cut the Harvard lead to 34-18.

The Crimson went nowhere on its next possession and a punt by senior Jake Dombrowski was blocked, giving the Tigers good field possession once again.

And once again Princeton took advantage, scoring on a 29-yard touchdown pass by Michelsen to Matt Costello. Epperly passed for another two-point conversion to cut the lead to 34-26.

The Crimson seemed ready to post a response on the final drive as it had a first-and-goal from the five-yard line. But the Tigers stuffed three straight Harvard runs and then blocked a field-goal attempt.

Michelsen then led Princeton 79 yards in just over two minutes to cut the lead to 34-32 before a third two-point conversation attempt was broken up.

The Crimson went three and out again on the following possession, but this time a Dombrowki punt pinned the Tigers at their own 10.

That set up a wild final possession for Princeton. Michelsen was hurried on first down by senior Nnamdi Obukwelu, leading to an incompletion. But then he completed a 15-yard pass, and then an eight-yard pass after another incompletion. A sack knocked Michelsen out of the game with less than a minute to go, but Epperly was effective after coming in, eventually heaving a pass into the back corner of the end zone that was brought down by Wilson.

"Roman [Wilson] is a great matchup and we needed to get the ball down the field so I took a shot," Epperly said. "And Roman came down with it."

Billed as one of the Crimson’s toughest tests of the year, the contest was a blowout early on as the visitors jumped out to a 20-0 lead.

On the second drive of the game, senior quarterback Colton Chapple found junior Ricky Zorn streaking down the sideline for a 52-yard score.

Attempting to respond, the Tigers went three-and-out on their next drive and senior D.J. Monroe blocked a punt attempt to give his offense the ball at Princeton’s 16-yard line.

One play later, the score was 14-0 thanks to a 16-yard pass from Chapple to Juszczyk. Chapple found Juszczyk again on the team’s next drive after two red-zone penalties had forced Harvard into a first-and-30 situation.

Yet Harvard had its extra-point attempt blocked after Juszczyk’s second touchdown, the first of several miscues that took points off the board for the visitor.

A fumble by senior running back Treavor Scales inside Princeton’s 30 and a Chapple interception in the end zone both ended scoring threats deep in Tiger territory.

"It’s a game where we could have scored 50," Murphy said. "We did some things really well, but we just had too many missed opportunities."

Princeton managed just 46 yards through the air between the two quarterbacks it rotated in and out in the first half, and the running game was just as ineffective as a slew of runners combined for just five yards at halftime.

—Staff writer Jacob D. H. Feldman can be reached at jacobfeldman@college.harvard.edu.