There was a six-man breakaway as the riders began climbing, with the peloton more than six minutes adrift, and Enric Mas went clear of Julian Alaphilippe and Thymen Arensman.
Vingegaard’s Visma-Lease a Bike team-mates took turns to push the pace in the peloton and, with Pogacar getting isolated from his team-mates early in the climb, Vingegaard launched three attacks, but each time the three-time Tour winner stayed on the Dane’s wheel.
Up the road, Healy and Paret-Peintre managed to catch Mas about 3.5km from the line and the trio got engaged in a tactical stalemate, allowing Santiago Buitrago to join them and set up a gripping final 2km.
After Mas faded, Ilan van Wilder suddenly charged into the lead inside the final kilometre and signalled for team-mate Paret-Peintre to follow.
And although Healy kicked first in the final 250m, Paret-Peintre had enough left in the tank to snatch victory.
Pogacar mounted one late attack and, although Vingegaard stuck to his wheel, the Slovenian then managed to sprint to the line to gain two seconds on the two-time Tour winner.
Merlier could go for a third stage win on Wednesday, as the race continues with a 160.4km flat stage from Bollene to Valence.
