We all know about the popularity of manga and anime - especially in Japan.

In the UK, Comic Con is the best place to find major expressions of manga fandom, but in the country from which it originates, the level of devotion is best summed up by the Japanese word 'otaku' - a term that has become a vogue in recent years to describe those who go beyond fanaticism.

Well, guess what? This obsession doesn't even appear to be restricted to humans. A penguin in Tobu Zoo in Saitama Prefecture has died, leaving behind the story of his long-standing love affair - with a cardboard cut-out of a cartoon character.

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Grape-Kun, a 20-year-old Humboldt penguin, fell in love with an advert of a manga character that was placed in his enclosure and spent his days staring intently at the image.

He had gradually spent less and less time with his fellow penguins after his long-term partner, another penguin called Midori, left him for a younger penguin. Yep, a classic tale of love and loss. Tragically, Grape-Kun had become increasing solitary - until the manga character was added to the penguin area at Tobu Zoo.

The image depicts a young girl by the name of Hululu, a well-known figure in Japan from the anime series Kemono Friends. Hmm, I suppose she does look a little bit like a penguin, wearing a white dress with black shoulders. No? Hey, it's the best I've got.

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Zoo staff spotted Grape-Kun performing courtship rituals to the sign, and when it was removed during a typhoon, the lovesick penguin was visibly distraught. Happily, staff returned it to its rightful place as soon as possible.

Following his death, Tobu Zoo tweeted "The Humboldt penguin Grape-kun passed away yesterday. Sincere thanks to everyone for supporting him until now. Thank you also to Hululu, who watched over him until the very end. And thank you, Grape-Kun, for all this time. Rest peacefully in heaven."

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The image of the penguin and his cardboard girlfriend was a viral hit around the world and the zoo regularly tweeted about the feathery romantic and his antics with the cartoon character.

Like many of the kids whose obsession with anime goes a little too far, Grape-Kun was also unable to turn his devotion into a meaningful relationship - although in his case it was because he couldn't fly up to the ledge where Hululu was standing.