An underwater photo of sardine migration on the Wild Coast of South Africa has been selected from thousands of entries as the grand prize winner of the 2016 National Geographic Nature Photographer of the Year contest. Greg Lecoeur from Nice, France, wins a 10-day trip for two to the Galápagos with National Geographic Expeditions and two 15-minute image portfolio reviews with National Geographic photo editors. He took the photo in June 2015 after waiting two weeks to witness the natural predation on sardines captured in the photo.

"During the sardine migration along the Wild Coast of South Africa, millions of sardines are preyed upon by marine predators such as dolphins, marine birds, sharks, whales, penguins, sailfishes and sea lions. The hunt begins with common dolphins that have developed special hunting techniques to create and drive bait balls to the surface," Lecouer said. "In recent years, probably due to overfishing and climate change, the annual sardine run has become more and more unpredictable."

2016 National Geographic Nature Photographer of the Year Contest
1st Place, Action and Grand Prize Winner: Greg Lecoeur, Sardine Run Greg Lecoeur/2016 National Geographic Nature Photographer of the Year Contest

Lecoeur's photo won the Action category. Varun Aditya, of Tamil Nadu, India, won the Animal Portraits category for a photo of a snake; Vadim Balakin, of Sverdlovsk, Russia, placed first in the Environmental Issues category for a photo of polar bear remains in Norway; and Jacob Kapetein of Gerland, Netherlands, placed first in the Landscape category for a photo of a small beech tree in a river.

IBTimes UK presents a gallery of the other images selected by the judges of the 2016 National Geographic Nature Photographer of the Year contest.