NASHVILLE (AP) — The Tennessee Titans' ninth and final penalty could not have come at a worse moment.

Overtime.

Trying to protect a 28-25 lead with the clock winding down, Kyle Peko sacked Colts quarterback Gardner Minshew on Indianapolis' third play for an 8-yard loss. A penalty flag thrown at Titans cornerback Sean Murphy-Bunting for illegal contact with a receiver wiped it out.

Minshew went right back at Murphy-Bunting and found Alec Pierce for a 55-yard pass down to the Tennessee 4. Two plays later, the Colts finished off a wild game with a 31-28 victory Sunday to cap a costly day for the Titans.

"We lost a tough game, hurt for the players because there were opportunities for us to give in, circumstances, situations," Titans coach Mike Vrabel said. "Guys were into it on the sidelines. Just unfortunate we didn't win."

The Titans lost their first home game in Nashville after being flagged nine times for 74 yards compared to just four for 29 for the Colts (7-5).

They also were swept by their AFC South rival for the first time since 2018 in a loss that may have extinguished their fading playoff hopes.

Three-time All-Pro wide receiver DeAndre Hopkins, still upset at a no-call that forced Tennessee to settle for a field goal at the end of the first quarter when targeted by Will Levis in the end zone, made his feelings over the officiating very clear, yelling at the nearest official.

Hopkins remained unhappy that Colts safety Nick Cross wasn't flagged for defensive pass interference especially after seeing what was called on his own teammate. The lack of a flag cost the Titans (4-8) four crucial points.

"He wasn't playing the football, and he was restricting me from running," Hopkins said of Cross. "So I don't know what a penalty is if that's not one."

Vrabel noted the flag on Murphy-Bunting was the first time the crew led by referee Craig Wrolstad had called illegal contact in the game.

The Titans led 17-7 and were dominating early and still took a 28-25 lead in overtime on a 46-yard field goal by Nick Folk. They held the ball for nearly 10 minutes more than Indianapolis and outgained the Colts 381-355 despite Levis being sacked six times.

But they finished without two-time Pro Bowl defensive lineman Jeffery Simmons, who left midway through the third quarter with an injured right knee.

Derrick Henry, the two-time NFL rushing champ, left early in the fourth after getting his third 100-yard rushing game this season to be evaluated for a concussion. He finished with 102 yards rushing and two touchdowns, tying him with Priest Holmes for 15th in NFL history with his 86th career rushing TD.

Punter Ryan Stonehouse left on crutches with his left leg in a brace. He came in leading the NFL in gross punting and was just selected as the AFC special teams player of the week. After having a punt late in the third quarter blocked and returned for a touchdown, Stonehouse had his next attempt blocked.

Tony Brown timed up the snap perfectly, sprinting away from the left before tipping the ball out of Stonehouse's hands and crashing into the punter's left leg.

That left Folk to punt twice. The veteran, who just turned 39 last month averaged 39 yards net. But he missed his first extra point this season working with backup quarterback and holder Ryan Tannehill, who filled in for Stonehouse as the usual holder.

"We just need to make sure that we protect our punter, protect our quarterback, and affect their quarterback, and affect their punter," Vrabel said. "That's what needs to happen."

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