The Southern defense had arguably its best game of the season Saturday, and so did the linebackers.

The Jaguars allowed just 220 total yards (their lowest total this season) and 99 rushing yards (their fewest against an NCAA opponent) in a 30-20 victory against Texas Southern. Southern allowed just 66 yards while shutting out the Tigers in the second half.

The performance featured three linebackers among the top four tacklers. Brian McCain had seven total tackles, including one sack and another tackle for loss. Demetrius Carter had six tackles, including half a tackle for loss. And Martin Henry had five tackles, including half a tackle for loss. Only safety Dionte McDuffy (nine tackles) had more tackles than the linebackers.

Of the 99 rushing yards, 34 came on one play when the defense “blew a blitz,” linebackers coach Lorick Atkinson said.

“Other than that, we held them pretty well intact,” Atkinson said before practice Monday. “(The linebackers) have elevated their game, and we’re seeing the results, especially in the run game. They are playing at the level we expect them to play at in November.”

Each month has brought changes to the linebacking corps. They learned in August that they would be without senior strongside linebacker Daniel Brown, a preseason All-Southwestern Athletic Conference second-team selection who was declared ineligible.

In September, McCain, who played linebacker in high school, volunteered to play defense. Later in the month, Javon Allen was lost for the season after hand surgery.

The Jaguars continued to tinker with the lineup as the weeks went by. Dorsey Bindon started in Brown’s spot in the opener, freshman Aaron Tiller started the next three games and then Roshaud Turner started four and Henry the past two.

“We had to move a lot of guys around, but we stayed the course,” Atkinson said.

Southern has settled on a starting group of Carter (the only linebacker to start all 10 games), Henry and McCain. Carter played 12 games and made 19 tackles as a sophomore last season.

“He plays the game the way it’s supposed to be played,” head coach Dawson Odums said. “He plays hard, he communicates, he gets our guys lined up.”

Turner, who had nine tackles in eight games as a freshman, was the only other linebacker on the two-deep to play with Southern in 2013.

“We’re being a lot more aggressive,” Atkinson said. “They’re a lot more comfortable with the scheme. Our main goal is to make sure they get lined up right and, when they get lined up, we let them go hunt.”

Carter, who leads the team with 83 tackles, said the defense plays better when it’s aggressive.

“That’s our identity,” he said. “When we lay a hat on people, we make big plays. We make wide receivers second-guess about catching balls, and it makes the running back’s speed slow down when he’s trying to hit that hole.”

Turner is fourth on the team with 41 tackles, Henry is fifth with 39 and McCain is eighth with 36.

Henry is a junior-college transfer who was recruited to play outside but was moved inside after Allen was injured.

“It’s taken him about eight weeks to get more comfortable with the scheme,” Atkinson said. “It’s a lot different between inside and outside.”

McCain had an even more challenging adjustment, but he moved into the starting lineup after four games and has started the past five, though he said he still needs an occasional tip from a teammate on his responsibility on a given play.

“It’s still taking some getting used to,” he said, “but we connect with each other, we motivate each other, we critique each other when we need to. I couldn’t ask for a much better group than the one I’m performing with.”

Now that Southern has settled on a starting group, the reserves — such as Tiller, fellow freshman Kentavious Preston and Turner — have become more comfortable with new roles.

Atkinson said Tiller’s “confidence is back up” after he struggled in his early stint as a starter.

“There’s no time for freshmen in November,” he said. “So Aaron has come off the edge for us and really played well. He’s growing up right in front of our eyes. Roshaud and Kentavious have come in and given us some good backup work.”

McCain said the enthusiasm of the younger players has been infectious as the Jaguars have won five in a row heading into their home finale against Mississippi Valley State on Saturday.

“You know how excited they are to win and how much they want to win,” McCain said. “That’s one thing that I believe makes our defense play the way it’s been playing.”

Odums said the linebackers are “really smart football players” who have “really relished the role of becoming very good linebackers.”

It has taken a while for the coaches to figure out how to best use the newcomers and for the players to grow into their evolving roles.

“You went from what on paper you thought was going to be a dominant group to, ‘You know what? Let’s see what they can do and put them in position to make plays,’ ” Odums said.

“That’s been the chess match, just trying to figure out what these guys can do very well and trying to put all those pieces together. It’s taken us a little longer, but I think we’ve stumbled onto something our guys can do.”