A ghostly shot of the flight paths of Daubenton’s bats, which took months to capture, has scooped top prize in this year’s British Wildlife Photography Awards.

Among the winning shots in the contest were the following:

A portrait of a badger

A great skua eating a puffin

A magpie on railings in the snow

A close-up of a nursery web spider waiting among petals for her prey

Grey seals being released

This shot of a nursery web spider won the Hidden Britain category Credit: Andrew McCarthy/BWPA/PA

Neil McIntyre won an award for the British Seasons category of Seasonal Scottish Squirrels. Credit: Neil McIntyre/BWPA

Paul Colley, from Swindon, Wiltshire, won the overall prize of £5,000 for his image Contrails At Dawn of Daubenton’s bats at Coate Water Country Park.

Capturing the flight of the high-speed mammals in the dark required an infrared camera and lighting system that was 14 months in development, and as the bats are a protected species they were photographed following advice from conservation experts.

David Bennett won an award in the Close To Nature category for his photograph of Goose Barnacles. Credit: David Bennett/BWPA

A picture of seasons overlapping in a beech wood won the Wild Woods category Credit: James Roddie/BWPA/PA

Mr Colley said: "No other image in my portfolio had been so clearly conceived and yet so difficult to achieve.

"My artistic intent was to capture this extraordinary little bat’s speed of movement and hunting flight path, but the journey to success was littered with disappointing failures."

He said he was supported by fellow photographers in his endeavour, which saw him experience a "huge gradient of emotion".

A Kelp Bed At Dawn won the Botanical Britain category Credit: Robert Canis/BWPA/PA

The winner of the 12-18 years category was Ivan Carter with a shot of tadpoles. Credit: Ivan Carter/BWPA/PA

Mr Colley added: "There were the lows felt during months of long, cold and exhausting dusk-to-dawn sessions, sometimes waist-deep in water and often without getting a single usable image.

"And then the natural highs of those lightbulb moments, when new ideas blossomed, problems were solved and the project inched closer towards the potential to win this exceptional accolade."

Shots of red squirrels won the British Seasons category. Credit: Neil Mcintyre/BWPA/PA

Prizes for top pictures were awarded in 15 categories, including ones that focus on the coasts, close-up images of the natural world, the same subject through the seasons, video, and a documentary series of photographs.

In the junior categories, Ivan Carter, 17, from Deal in Kent, won an award for his shot of common tadpoles, and nine-year-old Lucy Farrell, from Sunderland, Tyne and Wear, scooped top prize for a close-up of a cockchafer beetle.

Christopher Swan won an award for Urban Wildlife for his photograph of a Magpie in the Snow. Credit: Christopher Swan/BWPA

Lucy Farrell, aged nine, won one of the junior categories with her close-up shot of a cockchafer. Credit: Lucy Farrell/BWPA/PA

Naturalist, author and wildlife TV producer Stephen Moss said: "The extraordinary range of subjects, species and habitats, and the imaginative way they are portrayed, leaves us in no doubt that we in Britain are fortunate to be home to some of the most talented photographers in the world.

"It is also a snapshot of Britain’s diverse and beautiful wildlife, at a time when these wild creatures - and the places where they live – are under threat as never before."

Andrew Parkinson won an award in the Habitat category for his photograph of a Mountain hare. Credit: Andrew Parkinson/BWPA

Tesni Ward won an Animal Portraits award for her picture of Bean, a Badger. Credit: Tesni Ward/BWPA

An exhibition of winning and commended entries from the competition will go on tour, starting in London on November 6, and a book, British Wildlife Photography Awards 9, will feature the best images.

The category winners are: