ANNAPOLIS, Maryland - The popular African penguin known as Mr Greedy, a fixture of the Maryland Zoo who had sired many offspring and left “an astounding 230 descendants” over five generations, has died after an age-related decline in his health, the zoo said in a statement.

He was 33 – yes, in human years. (The zoo said it had no accurate way to determine the equivalent in penguin years.)

He was the oldest penguin in his colony, which the zoo said is the largest in North America.

When he was not busy reproducing or bringing joy to zoo visitors, Mr Greedy swam hard, took care of his mate – nicknamed Ms Greedy – and was constantly looking for things to steal.

His mischievous ability to steal nesting materials and food from others had earned him the affectionate nickname by which he was known, Ms Jen Kottyan, the bird curator at the zoo, said in a phone interview on Sept 7.

Mr Greedy, born in 1991, had a more official, though less personable name: African penguin No. 821. The cause of death was euthanasia on Aug 27, the zoo said.

In his long life, Mr Greedy “made a tremendous contribution to his endangered species,” said the statement from the Maryland Zoo, which is in Baltimore.

“This one bird was incredibly important to the continued existence of African penguins throughout the world,” Ms Kottyan said in the statement.

Mr Mike Evitts, a spokesman for the Maryland Zoo, said there is not a definitive number for the total descendants that are expected from a single African penguin, but that Mr Greedy’s tally is “way more than average.”

Found near islands off the south-western coast of Africa and on land near Cape Town, South Africa, African penguins can dive more than 122m underwater and can stay below the surface for two to...