Hundreds of puffins washed up on the east coast of the UK are likely to have died of starvation as a result of the recent severe weather.
RSPB Scotland said it has taken numerous calls from members of the public about the birds, found on beaches stretching from Aberdeen and Angus down to Northumberland.
It is the worst puffin "wreck" — the death of a large number of seabirds in a single incident — in almost 50 years, the conservation charity said.
Many razorbills and guillemots have also perished, prompting fears about the upcoming seabird breeding season.
Dr Barnaby Smith, from the Centre for Ecology & Hydrology which is part of the Natural Environment Research Council, said the birds may have been using up all their resources just fighting against the unseasonably cold temperatures and strong easterly winds.
This means they would have become exhausted and unable to find enough food to survive.
A RSPB Scotland spokeswoman said: "This may be the worst puffin wreck we have seen for almost half a century.
"Despite their small stature puffins are fairly hardy birds, adept at coping with the harsh conditions of life at sea. To hear that so many have been discovered dead is unusual and worrying.
"We are in close contact with experts from the Centre for Ecology & Hydrology to learn more about what is happening but it appears that the prolonged and unprecedented weather is making life extremely difficult for this species.
"We are fast approaching the start of the seabird breeding season where tens of thousands of seabirds return to their colonies to raise their young. The recent events could have an impact on the success of this year's puffin breeding season, a species already suffering population declines.
"RSPB Scotland, with the help of volunteers, will be closely monitoring the fortunes of this species and many other seabirds throughout the summer months."
Professor Mike Harris, also from the Centre for Ecology & Hydrology, has been keeping an online blog detailing the events.
"There is currently a major wreck under way which is the largest in the North Sea for at least 60 years.
"This was first noticed at the end of last week by people who regularly check the beaches of north-east England for dead birds. Then over the weekend reports started to come in of dead and dying puffins in beaches all the way north to Aberdeenshire.
"Most birds were emaciated and had obviously died of starvation.
"To date, I have heard of maybe 400 dead puffins and there will undoubtedly be many more, perhaps thousands, and this compares with just a handful over a whole typical winter. This will certainly be the largest wreck of puffins in the North Sea for over 60 years."
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