The puffin population of the North Sea coast has remained constant despite the death of thousands of the birds due to severe weather along the coast of eastern Scotland and north-east England at the start of spring.
Around 3,500 puffins were found dead along the coast in March, many of them breeding adults from local colonies.
But a count of the birds, funded by Scottish Natural Heritage, found that their numbers at Isle of May National Nature Reserve, on the edge of the Firth of Forth, are at similar levels to that recorded in 2009.
The survey by scientists from the Centre for Ecology & Hydrology indicates that a total of 46,000 burrows showed signs of use by puffins this spring.
The Isle of May reserve is home to the largest colony of puffins in the North Sea area and is a UK hub of research into puffins.
The count also revealed that the effects of the March weather seriously disrupted breeding on the island, with laying two to three weeks later than normal.
It is possible that some birds will not breed this year.