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 RunGreg 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.

2016 National Geographic Nature Photographer of the Year Contest
1st Place, Animal Portraits: Varun Aditya, Dragging you deep into the woods: 'I shot this at Amboli, Maharashtra, India, on 24 July 2016, during a morning stroll into the blissful rain forest. Ceaseless drizzles dampened the woods for 10 hours a day; the serene gloom kept me guessing if it was night or day. The heavy fog, chilling breeze, and perennial silence could calm roaring sprits. And there I saw this beauty. I wondered if I needed more reasons to capture the habitat, for I was blessed to see this at the place I was at. I immediately switched from the macro to the wide-angle lens and composed this frame.'Varun Aditya/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
1st Place, Environmental Issues: Vadim Balakin, Life and Death: 'These polar bear remains were discovered on one of the islands of northern Svalbard, Norway. We do not know whether the bear died from starving or ageing, but as we see the teeth are in good condition, it was probably from starving. They say nowadays that such remains are found very often, as global warming and the ice situation influence the polar bear population.'Vadim Balakin/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
1st Place, Landscape: Jacob Kaptein, Struggle of life: 'Last year I participated in the Marius van der Sandt Beurs. This scholarship stimulates photography by young photographers. For a whole year I was guided by some excellent nature photographers to realise a project I wanted to accomplish. I chose a natural stream restoration project of a nature organisation in the Netherlands. The first time I entered this patch of forest, I immediately saw this little beech. I came back several times to photograph it. One evening, just after sunset, all the light conditions were perfect. I stood in the cold water for more than an hour making many photos while I experimented with different shutter speeds.'Jacob Kaptein/2016 National Geographic Nature Photographer of the Year Contest

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

2016 National Geographic Nature Photographer of the Year Contest
2nd Place, Landscape: Alessandro Gruzza, Wild rink: 'The first cold days of winter have frozen the surface of a pond, and the first snowfall has revealed its delicate beauty. In low-pressure conditions, southwest winds push the clouds against the vertical peaks of the Pale di San Martino. At dusk, a long shutter speed enhances the movement of the clouds around Cimon della Pala, one of the highest peaks in the Dolomites.'Alessandro Gruzza/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
3rd Place, Landscape: Santiago Borja, Pacific Storm: 'An isolated cumulonimbus storm developed over the Pacific Ocean a few miles south of the coast of Panama City. It sat atop a temperature inversion that created a thick overcast layer of clouds. The strong updrafts of the storm quickly reached the tropopause and spread out, creating the characteristic anvil. The strongest updrafts pierced the tropopause and turned into what scientists call the overshooting tops. The entire frame was lit by a single lightning from within the storm in a moonless night on 16 June 2016.'Santiago Borja/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Honourable Mention, Landscape: Prasenjeet Yadav, Serendipitous Green Meteor: 'Anand Varma was visiting me and I was showing him around a mountain range in South India called the Western Ghats. We camped on the side of a road and I set up my Nikon D600 and a 24-70mm lens to take 15-second exposures. I set the camera to take 999 images. I slept next to the camera and it continued taking pictures until dawn. It wasn't until the next afternoon that I reviewed my images and noticed something unusually bright and green. I showed it to Anand, and we realised that I had captured an extremely rare event. After checking with a few experts, I learned that it was a green meteorite, and getting it on camera is very rare. This is an example of being at the right place at the right time to capture something totally unexpected. For those 15 seconds, I was the luckiest photographer on the planet.'Prasenjeet Yadav/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
2nd Place, Environmental Issues: Chris McCann, Outside Facebook HQ: 'Eighty percent of the San Francisco Bay Area wetlands – 16,500 acres – has been developed for salt mining. Water is channelled into these large ponds, leaves through evaporation, and the salt is then collected. The tint of each pond is an indication of its salinity. Microorganisms inside the pond change colour according to the salinity of its environment. This high-salinity salt pond is located right next to Facebook headquarters, where about 4,000 people work every day.'Chris McCann/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
3rd Place, Environmental Issues: Eleanor Ryder, Toxic Vanity: 'This image is a magnification of plastic particles in eyeliner, exploring just one facet of the synthetic swarm suspended in our oceans. The particles, lash-lengthening fibres, illuminating powders, and glitters these products contain are in fact tiny pieces of plastic. Every time we wash these products from our bodies or ingest them as we lick the glosses from our lips, we unknowingly add to the trillions of microplastic particles currently infesting every level of the ocean. This photograph was taken at Falmouth University in Cornwall, United Kingdom, in May 2016 using a reflective photographic microscope with a stacking panoramic process to create the final image.'Eleanor Ryder/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Honourable Mention, Environmental Issues: Patty Waymire, No snow, no ice? 'A solitary bear sits on the edge of Barter Island, Alaska. There is no snow when, at this time of year, there should be. The locals in Kaktovik noted that it's been an unseasonably warm winter, and that the ice will be late in forming this year. This will have an impact on the local polar bear population when it comes time to hunt seals for their food in the winter months.'Patty Waymire/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Honourable Mention, Environmental Issues: Ken Bower, American Flowers #1: 'In East Greenland's landscape lies Bluie East Two, a remote US Air Force base from World War II. The base was abandoned in 1947 and everything was left behind: military vehicles, structures, 800-plus cases of dynamite and munitions, and over 10,000 aviation fuel barrels. The Inuits who live in the region call these rusted remains American Flowers. I made it to the base in 2014, but I had to return to photograph it again in 2015. I wanted to get there early in the season when there would still be snow on the ground to provide contrast.'Ken Bower/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Honourable Mention, Environmental Issues: Sergej Chursyn, Wildfire at the beach: 'A young woman in a bikini looks at an approaching forest fire near the beach. A firefighting plane drops water to extinguish the wildfire. This image was taken at the beach of Son Serra, on the island of Mallorca, on 18 August 2016.'Sergej Chursyn/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
2nd Place, Animal Portraits: Michael O’Neill, Proud Momma: 'A female peacock bass guards her brood in a Miami, Florida, freshwater lake. She will protect her young fry from a variety of predatory fish until they are large enough to fend for themselves. This tropical freshwater species, also known as the peacock cichlid, was introduced in Florida in the mid-1980s from South America to control the tilapia population, another invasive species. Throughout its native range (and in Florida) it’s a prized sport fish known for its fighting spirit.'Michael O’Neill/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Equal 3rd Place, Animal Portraits: Mario Suarez Porras, Puffin Studio: 'This image was taken during the summer of 2015 on Skomer Island, Wales. This island is well known for its wildlife and the puffin colony is one of the largest in the UK. The photo shows a detail or study of an Atlantic puffin resting peacefully under the rain. As Skomer is not inhabited, puffins do not feel afraid of humans, and people can get really close to puffins. That morning, the conditions were perfect. Both fine rain and a soft light, so much appreciated by photographers, helped to take this picture. In order to get this angle from above the bird, I couldn’t make use of the tripod, as it could disturb the puffin. The photo had to be taken handheld, which added an extra challenge.'Mario Suarez Porras/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Equal 3rd Place, Animal Portraits: Jose Pesquero Gomez, Friendship knows no colour: 'Two Empusa pennata on the same plant was the rare scene I found when I visited one of my favourite locations for macro shots. This area is located near a village called Las Rozas in Madrid, Spain. There’s a small stream about one kilometre long where you can find a varied ecosystem with many different types of insects and arachnids. From May to September, I had seen up to four different Empusas alone on their plants, but on this day I was extremely lucky when I found two individuals on the same plant. I took advantage of such a discovery and mounted my macro set and took several photos of this magical scene, where the Empusas seem to play or dance, sharing the same plant like good friends.'Jose Pesquero Gomez/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Honorable Mention, Animal Portraits: Lawrence Chia Boon Oo, Crow chasing Buffy Owl: 'This shot was taken on an early September afternoon along the riverbank at Pasir Ris Park in east Singapore. I was hoping to capture wildlife in action when some movement in the bushes nearby caught my attention. Instinctively, I prepared my equipment to capture any action that might ensue. I was fortunate enough to witness this adult crow chasing an adult buffy fish owl right in front of me, proving at once that the crow was the more aggressive species of the two. The entire spectacle between these day and night creatures lasted less than two seconds and exemplified nature in its uninhibited form.'Lawrence Chia Boon Oo/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
2nd Place, Action: Tori Shea-Ostberg, Approach: 'An EF2 tornado bears down on a home in Wray, Colorado, on 7 May 2016. As soon as we were safe, as the tornado roared off into the distance through a field before roping out, we scrambled up the hill to check on the residents. Thankfully, everyone was all right, and we were grateful for that. As I was checking in with a young woman coming out of the basement, we became very aware of a strong new circulation right above our heads. We needed to run for cover and did so before saying a proper goodbye.'Tori Shea-Ostberg/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
3rd Place, Action: Zsolt Kudich, Great Egrets Take Flight: 'A remarkable conservation success story, the graceful great egret was saved from the brink of disappearance in Hungary, where in 1921 there were only 31 mating pairs remaining. Less than a century later, international conservation efforts have triumphed. We can now count over 3,000 mating pairs in Hungary alone. Today, although their numbers are continuing to climb, the great egret remains the Hungarian Nature Conservation’s symbol, and they are still considered an endangered species that must be carefully protected. The birds are most active at dawn and dusk, and here they’re squabbling over food and jostling for space in the crowded swamp.'Zsolt Kudich/2016 National Geographic Nature Photographer of the Year Contest
2016 National Geographic Nature Photographer of the Year Contest
Honourable Mention, Action: Scott Portelli, Jellyfish Feast: 'I came across this jellyfish on a dive in Byron Bay, Australia. At the time there was a single turtle eating various parts of the jellyfish. Soon, two more turtles arrived in full force and started a tug-of-war to keep the best bits of the jellyfish. I moved in closer to position myself in between the action as the turtles snapped at each other and continued to eat, oblivious to my presence. Soon there was little left of this giant jellyfish, and a small group of fish were evicted from their shelter within the stinging tentacles.'Scott Portelli/2016 National Geographic Nature Photographer of the Year Contest