Jahwan Edwards broke a tied game with a 15-yard touchdown run late in the fourth quarter, as Ball State ended Toledo's eight- game winning streak by topping the 23rd-ranked Rockets, 34-27, at the Glass Bowl.
Edwards ran for 105 yards on 16 carries, and Keith Wenning threw for three touchdowns for the Cardinals (7-3, 4-2 MAC), winners of four straight following losses to Kent State and Northern Illinois.
Wenning completed 29-of-42 passes for 280 yards and was intercepted three times, but Ball State made up for the miscues by forcing a key turnover on downs midway through the fourth quarter.
"It doesn't get any better than this," Ball State coach Pete Lembo said. "Our guys just hang in there and keep battling. We're not a great team but we're a team. We're together. We ride out the highs and lows and we keep believing."
Toledo's David Fluellen gashed the Cardinals for 200 yards and a score on 34 carries, though he lost a fumble and was unable to move the chains at a critical time.
Terrance Owens threw for 215 yards and a TD on 14-of-27 throws for the Rockets (8-2, 5-1), who had not lost since dropping their season opener to Arizona in overtime.
"Ball State made the more disciplined plays throughout the football game," Toledo coach Matt Campbell said. "We played behind much of the game tonight, and we did some things that have been uncharacteristic of us this season. We have to correct those and pay attention to details and executing in critical situations as we move forward. Hopefully tonight is a great learning experience."
With the score tied, 27-27, Campbell opted to pass up a short field goal try. It was the second time in the fourth quarter Toledo's head coach went for the first down on 4th-and-1. The first gamble resulted in a touchdown run by Fluellen, but the second ended with Fluellen getting stuffed for no gain at the Ball State 14-yard line with 6:02 remaining.
Edwards did most of the work in getting the Cardinals down the field, carrying the ball four times in the first five plays for 41 yards, including a 19-yard run on 3rd-and-9. Another third-down conversion on a nice catch by Zane Fakes kept the drive alive, and Edwards took a handoff around the left side and sprinted into the end zone with 1:43 remaining.
Toledo, with no timeouts, only moved the ball seven yards on its final possession. A sack on second down and two straight incompletions sealed the upset win for the visitors.
Wenning went 7-for-8 for 71 yards on the game's opening drive and connected with Jamill Smith on a corner route in the end zone for a 6-yard score.
After Toledo punted in enemy territory, Wenning completed all five of his attempts on an 87-yard march, the last 25 yards coming on Connor Ryan's touchdown catch for a 14-0 lead less than 11 minutes in.
Later in the first quarter, Fluellen got Toledo moving with a 31-yard run, and Owens found an open Bernard Reedy behind the secondary for a 24-yard TD.
An illegal block negated a Fluellen touchdown run early in the second quarter, and the Rockets settled for a 29-yard field goal by Jeremiah Detmer.
Another promising drive by the Rockets ended when Fluellen fumbled near the red zone. Ball State linebacker Tony Martin scooped up the loose ball, ran it back to the Toledo 12, and the Cardinals padded their lead on Steven Schott's 24-yard field goal with just over four minutes left in the half.
Fluellen shook off the fumble and helped Toledo reach the Ball State 33, and Cassius McDowell picked up the rest on three carries to make it a 17-17 game heading to halftime.
The Rockets went three-and-out on their first two possessions of the second half, allowing Ball State to regain the lead. A pair of third-down conversions on a 10-play, 61-yard drive ended with Smith hauled in a 3-yard TD pass.
Detmer's 22-yard field goal cut Toledo's deficit to 24-20 late in the third quarter, and Dan Molls came away with an interception on the first play of the ensuing drive.
The Rockets looked like they were going to settle for three again when Owens was stopped a yard short on third down, but Campbell kept his offense on the field. Fluellen then took a handoff up the middle and went virtually untouched for a go-ahead 16-yard score on the first play of the fourth quarter.
Schott, who missed a 35-yarder midway through the third quarter, tied things again on a 30-yard field goal with 9:41 remaining.
Toledo, which was ranked for the first time since 2001, trails Northern Illinois by one game in the West Division. The two schools play each other next Wednesday ... Ball State went 8-for-12 on third-down conversions, compared to 7-for-16 for Toledo ... The Rockets outgained the conference's best offense, 526-447 ... Toledo committed 11 penalties for 115 yards ... Toledo holds a narrow 19-18-1 advantage in the all-time series.