Joe Paterno, major-college football’s all-time wins leader, died Sunday, according to a statement released by his family. He was 85. He had been undergoing treatment for lung cancer.

Joe Paterno, the winningest coach in college football history, has died at age 85. The former Penn State coach had struggled with lung cancer and was dismissed recently amid a sex-abuse scandal at the school. WSJ’s Darren Everson reports. (Photo: AP)

The man known by his Pennsylvania State University fans as JoePa will be remembered both for his legendary career leading one of his sport’s top programs and for the abrupt way that career ended in November. Mr. Paterno was fired by Penn State’s board of regents as part of the fallout from the arrest of former longtime Nittany Lions assistant coach Jerry Sandusky.

Mr. Paterno’s 46 years in charge of Penn State’s program, starting in 1966, earned him 409 victories, a pair of national titles, a statue in front of his team’s home field and a spot in the College Football Hall of Fame. His ties to the school extended to 1950, when he started as an assistant there.

During his time in State College, Pa., he and his wife Sue donated an estimated $4 million to the university. His Nittany Lions were also held up as a rare marquee football program that won without ever having been found guilty of major violations by the National Collegiate Athletic Association, college sports’ governing body. (The organization defines “major” violations as those involving acts that give a team a competitive or recruiting advantage.)

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Associated PressJoe Paterno died Sunday at 85.

The story of Mr. Paterno’s fast, involuntary exit from the school could not pose a starker contrast with his decades of success there. Mr. Sandusky, a longtime assistant of Mr. Paterno’s, faces numerous felony counts of sexually abusing 10 boys over more than a decade. Mr. Sandusky has maintained that he is innocent of the charges.

Mr. Paterno was ousted because of the way he responded to learning of an alleged incident involving Mr. Sandusky and a child in 2002. Upon hearing of the incident from an eyewitness, Mike McQueary (then a young assistant on the Penn State staff), Mr. Paterno reported it to school officials but not to police, according to a report issued by the grand jury. Although Mr. Paterno was not charged in the case, he quickly came under widespread criticism for not doing more, though some prominent alumni have spoken out publicly against his abrupt dismissal.

In a recent interview with the Washington Post, Mr. Paterno said that if Mr. Sandusky is guilty, “I’m sick about it,” and that he didn’t know how to handle the situation when it was brought to him. “I was afraid to do something that might jeopardize what the university procedure was,” he said. “So I backed away and turned it over to some other people, people I thought would have a little more expertise than I did. It didn’t work out that way.”

For years, college football fans wondered how long Mr. Paterno would coach. Until the Sandusky case broke, Mr. Paterno still hadn’t given any indication publicly when he would step down. As it turned out, because of his abrupt departure he never got an on-field send-off like former Florida State University coach Bobby Bowden and other top coaches. The Big Ten Conference—to which Penn State belongs—removed Mr. Paterno’s name from its championship-game trophy in the wake of the scandal.

Born Dec. 21, 1926, Mr. Paterno, who was from Brooklyn, N.Y., went on to attend Brown University, where he played quarterback under coach Rip Engle. When Mr. Engle became coach at Penn State in 1950, Mr. Paterno—who graduated that year—joined him.

In those days, Penn State was a football backwater. The school, located in the sparsely populated center of the state, had never won a football national title and had played in only two postseason bowl games.

But the program improved under Mr. Engle and took off under Mr. Paterno, who became head coach in 1966. He led the Nittany Lions to an 11-0 record in his third season, its first football national championship in 1982 and another national title in 1986—defeating favored Miami in a famous No. 1-versus-No. 2 battle in the Fiesta Bowl.

In the process, Mr. Paterno turned Penn State into one of the best-known brands in American sports. Dressed for decades in basic blue-and-white uniforms with no names on the backs of their jerseys, the Nittany Lions became synonymous with old-school, fundamental football. Beaver Stadium, Penn State’s home field, swelled to a seating capacity of over 100,000. Penn State began playing football in the Big Ten Conference in 1993, and Mr. Paterno led the Lions to a perfect season a year later.

Mr. Paterno’s Lions also became regarded as “Linebacker U” for their defensive prowess and their success in placing linebackers in the National Football League—more than 50 in all. One of the individuals most responsible for Penn State’s defensive reputation was Mr. Sandusky, who was an assistant coach—and later the defensive coordinator—from 1969 to 1999. He was once thought to be a potential successor to Mr. Paterno in the 1990s, but according to the grand-jury report in the Sandusky case, Mr. Paterno told Mr. Sandusky in a May 1999 meeting that he wouldn’t be the next head coach at Penn State. Mr. Sandusky retired following that season.

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Associated PressJoe Paterno died Sunday. He was 85.

In recent years, Mr. Paterno had largely become a figurehead as Penn State’s coach. He didn’t do the most visible coaching tasks nearly as much as other coaches typically do, like calling plays from the sideline or going on recruiting trips. Late in life, injuries repeatedly forced him to coach from the press box instead of the sideline during games.

The Penn State program also developed a reputation as being unusually insular. For years, Mr. Paterno’s salary—unlike almost all coaches at public universities—was unknown to the public. A change in state law finally forced disclosure in 2009. (According to filings the university has made public, Mr. Paterno made $1.02 million for the July 2009-June 2010 fiscal year.)

Before the Sandusky case, Mr. Paterno’s job was in jeopardy once before, in 2004. At that time, the Nittany Lions had endured four losing seasons in five years, their worst run in Mr. Paterno’s tenure. Mr. Paterno told the Pittsburgh Post-Gazette a year later that school administrators met with him and suggested he quit.

But Mr. Paterno rallied. The Nittany Lions finished the 2005 season ranked No. 3 in the country. This year’s team was also enjoying a surprisingly strong season, winning eight of its first nine games before the Sandusky case broke. It finished 9-4, with interim coach Tom Bradley in charge for the last four games. Penn State has since hired New England Patriots offensive coordinator Bill O’Brien as head coach.

Mr. Paterno’s death occurred less than three months after his final game as coach, and thus echoed the death of another longtime college football coach. Paul “Bear” Bryant, who led the University of Alabama to six national titles in the 1960s and 1970s, died in 1983 at 79, about a month after his last game. Both Messrs. Bryant and Paterno coached their teams to victory over the University of Illinois in their final game.

Write to Darren Everson at darren.everson@wsj.com

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