Bronson Yoder ran for 106 yards and a touchdown and the William and Mary defense picked off four passes, leading the Tribe to a surprising 31-18 over Villanova on the Main Line on Saturday.

The Wildcats, ranked fourth in the FCS coaches poll, were held well below their 34.6 scoring average and allowed more than double their 15.4 points-against average — both numbers were second in the Colonial Athletic Association.

Behind Yoder, who scored on a direct snap, the Tribe (6-2, 4-1 CAA) had 258 yards on the ground. Malachi Imoh, quarterback Darius Wilson, and JT Mayo all scored rushing touchdowns.

Villanova quarterback Daniel Smith completed 18 of 38 passes for 252 yards and a touchdown and ran for another score, but he threw all four interceptions.

The Wildcats (6-2, 4-1), whose only other loss was to Penn State, had 291 yards on offense, but 154 came on two fourth-quarter scoring drives after William and Mary took a 31-3 lead.