Red squirrels have returned to Perthshire and Fife for the first time in five years.
A survey by the National Trust for Scotland (NTS) found red squirrels at 29 trust properties including Branklyn Garden in Perth and Falkland Palace in Fife.
Experts described it as “good news”, and said it is evidence that work to control the non-native grey squirrel population in the areas is working.
The survey found there were red squirrels at 29 trust properties, and grey squirrels at 32.
NTS nature adviser Lindsay Mackinlay said: “The recent census of our properties has shown that red squirrels are holding their own, and even thriving in many cases.
“We’ve had some real successes in our Aberdeenshire, Perthshire and Fife properties, where we have seen the near disappearance of grey squirrels from places like Crathes and Drum after years of seeing them expand in numbers, whilst we have seen reds return in other places.
“Greys have expanded their range in some regions, particularly around House of Dun, beside Montrose, and they continue to threaten reds in Dumfries and Galloway.
“The current situation is stark and simple – greys are still here, and with squirrelpox virus moving northwards with them, there is a very real danger for our red squirrels in some of our most beautiful properties, like Killiecrankie, Crathes, Threave and House of Dun, to name but a few. All these properties sit on the so-called front line of grey expansion.”
The trust is planning to introduce red squirrels in the Highlands to protect numbers.
Mr Mackinlay added: “We are looking at our properties in the north west to see if they would be suitable for red squirrel introductions, and would encourage other landowners to do the same.
“This could provide a long-term refuge for red squirrels should grey squirrels and the squirrelpox virus keep heading northwards.”