LONDON — Scientists are 99.999 percent sure, in their most conservative estimate, that remains found in 2012 belong to King Richard III. These results, published Tuesday in Nature Communications, put a 529-year-old cold case to rest – all thanks to some intense genetic detective work.

The British royal died in battle in 1485. But while he would be immortalized in his namesake Shakespeare play, King Richard III was buried without fanfare. The church that marked his grave had long since been demolished when researchers went looking for it in 2012, and pinpointing its former location took some investigating. But when they finally tracked down Greyfriars Church, its foundation – and by extension, the body of a king – turned out to be under a modern-day parking lot.

Richard III’s skeleton was exhumed soon after and tentatively identified based on its age, battle wounds and signs of scoliosis – from which the king is believed to have suffered, based on historical descriptions. But to get a more certain identification, researchers needed to look at the skeleton’s DNA.

“Ancient DNA is far more difficult to work with,” lead study author Turi King, a geneticist at the University of Leicester, said.

King and her colleagues tracked down living relatives of the king and analyzed their genetic makeup, checking up and down the lines of inheritance to see whether the centuries-old skeleton fit into the family tree.

The combination of circumstances – someone with this particular DNA, who showed signs of scoliosis and died in battle at the same time and location as the king – means that there’s less than a one in 100,000 chance that the bones belong to someone else.

“The evidence is overwhelming,” King said.