A 'heartbreakingly poignant' image of a female elephant carrying a dead calf has won this year's Wildlife Photographer of the Year competition.
Italian photographer Federico Veronesi captured the black and white image after catching up with a family group of elephants in Namibia's Huab River Valley.
It was chosen from a record-breaking 65,403 entries from 121 countries and territories.
The calf is wedged between the elephant's tusk and trunk. Such behaviour around death has been observed in both African and Asian elephants.
Experts suggest it could reflect attempts to understand death or possible displays of grief.
Kathy Moran, chair of the Wildlife Photographer of the Year jury, said the winning photograph 'says it all'.
'This photograph works on so many levels,' she said.
'Emotionally it's a gut punch, heartbreakingly poignant. Intellectually it's proof that animals have cognition.'
A black and white shot – of a dung beetle pushing a ball of dung – also scooped the top prize for the Young Wildlife Photographer of the Year competition, which was won by Oscar Whitfield, a young British photographer living in Zimbabwe who was 14 when he took the picture.
Jasper Doest, jury member for the 62nd Wildlife Photographer of the Year competition, said the image was 'exceptionally well done'.
A 'heartbreakingly poignant' image of a female elephant carrying a dead calf has won this year's Wildlife Photographer of the Year competition
A young bull elephant spraying a group of zebra at a waterhole in Kenya, by James Lewin. This picture won the Animals in their Environment category
A dung beetle pushing a ball of dung, by Oscar Whitfield, from United Kingdom but living in Zimbabwe, the winner of Young Wildlife Photographer of the Year
A dead puma amid the rugged, windswept beauty of Argentine Patagonia, by Francisco Negroni, from Chile. The picture won the Photojournalism category
A common water monitor carrying its fish supper in a Bangkok park, by Seth Forestson, from United Kingdom. The picture won the Urban Wildfire category
a flesh fly with its needlelike mouthparts, by Alexis Tinker-Tsavalas, from Germany. This photo won the Rising Star Award category
Trees still swathed in autumn foliage, by Michal Michalski, from Poland. This photo is the winner of Natural Artistry category
A newt laying its eggs as it clasps vegetation that already holds another newt's egg clutch. The photo is the winner, of Behaviour: Invertebrates by Anton Sorokin, from United States of America
A scorpion burrowing into one of the shifting dunes of the Thar desert Sitaram Raul
A whale shark investigating the catch of a 'bagan', or mobile net-fishing platform, in the inky waters off Papua, by Alex Lindbloom, from United States of America, the winner of Oceans: The Bigger Picture category in the Wildlife Photographer of the Year
Shoebills gathering nesting material at Mabamba Swamps in Uganda, by Isak Pretorius, from South Africa, the winner of Behaviour: Birds category in Wildlife Photographer of the Year
A coyote pads along a trail in Bernal Heights Park, overlooking the city and bay of San Francisco by Loren Elliott, USA, the Winner, Photojournalist Story Award
A common carder bee coming into land on a flower, by Amelie Davis, from United Kingdom, the winner of 10 years and under category in Wildlife Photographer of the Year
A common octopus peeking out from kelp in the waters of southwest England, by Aaron Sanders, from United Kingdom, the winner of Animal Portraits category
A scorpion burrowing into one of the shifting dunes of the Thar Desert, by Sitaram Raul, from India, the winner of Winner, Behaviour: Invertebrates category
Fluorescent fungi, by Agorastos Papatsanis, from Greece, the winner of Plants and Fungi category
Slime moulds in the woodlands in West Sussex, by Kutub Uddin, from Bangladesh, the winner of Urban Wildfire category in the Wildlife Photographer of the Year
He added: 'Photographing such a small, constantly moving subject is challenging, yet the image feels so intentional.
'The composition reveals an entire story and by doing so the photograph quietly reveals how life cycles depend on what we often overlook or dismiss.'
The winning pictures, along with category winners from topics ranging from urban wildlife to animals in their environment, will feature in an exhibition at the Natural History Museum, London, which runs the competition each year, before a UK and international tour.
Among the category winners are fluorescent images of fungi in Greece, a common octopus peeking out from kelp in the waters of south-west England, a pair of elusive shoebills gathering nesting material in South Africa, and a young bull elephant spraying a group of zebra at a waterhole.
Photographers winning for a portfolio of images include Kutub Uddin, for his shots of the tiny hidden world of slime moulds in his local woodlands in West Sussex, England.
Young winners also included Amelie Davis, who scooped the top prize in the 10 years and under category for her shot taken when she was nine years old of a common carder bee coming into land on a flower, captured on her father's smartphone.
Dr Doug Gurr, director of the Natural History Museum, said: 'This year's winning photographs demonstrate the extraordinary power of wildlife photography to help us see nature differently.
'From a deeply affecting image of an adult elephant carrying a dead calf to the determination of a dung beetle rolling a ball many times its own size, these photographs capture moments that are at once intimate, surprising and profoundly revealing.
'Together they remind us that every species plays a role in the complex web of life, and why understanding and protecting the natural world has never been more important.'