Beavers have returned to the Cairngorms 400 years after the species were driven to extinction in Scotland.
A pair were released at Rothiemurchus Estate, near Aviemore, and another pair on land owned by billionaires Anne and Anders Holch Povlsen.
Over the next five years, 15 families could be released in the Cairngorms National Park.
Some landowners have raised concerns about damage the animals could do to farm and wood land.
The four beavers were released on Monday under a licence from public agency NatureScot.
The beavers had earlier been trapped in the River Tay’s catchment area before being taken to Five Sisters Zoo, near West Calder, West Lothian, for health checks.
The reintroduction project partners include Cairngorm National Park Authority, RSPB Scotland and the Polvsens’ Wildland Cairngorms Ltd, which manages the Gaick, Glenfeshie and Kinrara estates.
The upper River Spey catchment in the Cairngorms represents the fifth catchment to which beavers have either been officially granted permission to remain, or have been released.
Populations are already established in Tayside, on the Forth, in Knapdale and Loch Lomond.
Biodiversity Minister Lorna Slater assisted with the release of the beavers at the Rothiemurchus site.
She said: “Today is a key milestone in our ongoing collective efforts to ensure this iconic species can once again thrive across Scotland.
“I want to thank the Cairngorms National Park Authority and all the organisations involved who have made this truly exciting moment possible.”
Beavers died out in Scotland about 400 years ago but were reintroduced in 2009.
Conservation organisations say beaver activity can improve water quality and natural habitats and also help to alleviate flooding.
But some farmers have said a proper consultation was not held ahead of the Cairngorms releases.
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