BOSTON - A 22-game winning steak? No problem. On the road? No problem. Ivy League title at stake? No problem.

Penn handed Harvard its first loss since 2013 on Saturday, upsetting the Crimson, 35-25, in a showdown atop the Ivy standings. With the victory, the Quakers (6-3, 5-1 Ivy) have a chance to claim at least a share of the Ancient Eight title with a win over 1-8 Cornell next weekend.

"I'm very proud of this team and all we've accomplished," coach Ray Priore said. "But we have some unfinished business to do."

Penn came out of the gate quickly, going up, 14-0, in the first 10 minutes on a pair of rushing touchdowns by junior quarterback Alek Torgersen. The teams traded touchdowns, leaving Penn with a 21-6 lead after one quarter.

The second quarter belonged to the Crimson (8-1, 5-1), ranked 12th in the Football Championship Subdivision, as Harvard scored three touchdowns. Brothers Semart and Seitu Smith led the way, scoring within four minutes of each other midway through the period to give the Crimson a 25-21 halftime advantage.

Penn, however, did not allow another point. The Quakers excelled in containing Harvard standout Paul Stanton, limiting the Crimson's leading rusher to just 78 yards on 13 carries. Harvard also struggled on special teams, failing on three point-after plays and missing a 27-yard field-goal attempt.

Meanwhile, Penn's offense posted 14 unanswered points in the second half. Sophomore wideout Justin Watson provided the knockout punch, rushing for a 79-yard score with 13 minutes left to play - his second touchdown of the day - giving the Quakers a 10-point advantage they did not relinquish.

"I just had to get to the sideline and run straight. I've never seen a hole that big," said Watson, who had game highs in both rushing (100) and receiving (149) yards. "Those guys really opened it up. It's all them."

Penn, Dartmouth, and Harvard are 5-1 in league play, so wins by each squad next Saturday - Dartmouth hosts 5-4 Princeton, Harvard plays at 6-3 Yale - would result in a three-way tie for the conference crown.

Penn, which was picked to finish sixth in the Ivy League in the preseason coaches' poll, is undoubtedly the biggest eye-opener of the group.

And for the Quakers, Saturday's win was perhaps the biggest surprise yet.