Nine in ten of the 3,200 animals that died in the Oostvaardersplassen nature reserve during the winter were shot by rangers to control numbers, according to official figures.

Around 60% of the total herd of red deer, cattle and horses perished in the unusually long and wet winter spell, which concluded with a week of freezing temperatures at the start of March.

The forestry service Staatsbosbeheer said 2,684 red deer, 75 cattle and 467 horses died in the five months from December to April, NOS reported. Of these, 89% were culled because rangers decided they were not strong enough to survive the winter.

Last month an official inquiry into the nature reserve recommended reducing the size of the herd further to around 1,100 by the end of the autumn. The population had grown to more than 5,230 animals by October last year following a series of mild winters.

The fate of the animals led to heated protests and stand-offs between rangers and animal rights activists who tried to break into the reserve with food for the starving animals. The provincial administration in Flevoland recently agreed to extend the winter feeding season until May 5 in response to protesters’ concerns.