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.
"This March's wreck has clearly had a serious effect on the puffins on the Isle of May but, perhaps surprisingly, numbers are very similar to the last count which took place in 2009. "Our general impression over the last few years was that the population was increasing slowly and this may explain why we have not seen a decline following the recent wreck."
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.
"The March wreck has seriously affected the timing of breeding with those birds that did survive breeding very late. It would not be surprising if they needed a few weeks to recover and get into breeding condition. "We now wait to see how successful these birds are in raising chicks this summer."