Mr. Greedy, who helped keep his species of African penguin alive, died peacefully at his home on Penguin Coast.

Hatched in 1991, he moved to Baltimore as an adult a year later and took up residence at the Maryland Zoo. He is survived by his longtime companion, Mrs. Greedy, and 230 descendants.

Mrs. Greedy — also hatched in 1991, and who used to go by her maiden name, Penguin #832 — had also just arrived in the city. They engaged in a slow-burn romance, not pairing up until two years later. They have been inseparable ever since.

Wild penguins typically mate for life. In captivity, though, they can have multiple partners. Mr. Greedy was part of a program called Species Survival Plan, where penguins are matched up based on their genes and the needs of the colony’s population, based on how many offspring survive from each couple and their genetic lines. The aim is to ensure African penguins do not become extinct. He was matched with Mrs. Greedy based on their genetics and were only supposed to be together a few years.

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Usually, the zoo runs a computer program that determines who should be with whom to get the healthiest and most genetically diverse chicks out of the group, said the Maryland Zoo’s Bird Curator, Jen Kottyan.

Mr. Greedy was 33, the oldest penguin at the Zoo’s colony and “incredibly important to the continued existence of African penguins throughout the world,” Kottyan said. He died Aug. 27.

Mr. and Mrs. Greedy proved to be “genetically valuable,” Kottyan said. They reared many successful “clutches of offspring,” she said, so it made more sense to keep them together rather than separate them.

In other words, they were meant to be.

Penguin #821, aka Mr. Greedy, is in the center of this photo with a purple, brown, and black ID band. (Courtesy of the Maryland Zoo)

Mr. Greedy was protective of Mrs. Greedy, standing guard on the nest box when she was incubating their eggs. They would swap on and off every couple of days until the eggs hatched, and he made sure the chicks were being brooded, were warm and fed appropriately. His offspring were never underfed or unattended, Kottyan said, and Mr. Greedy was a good bird to be around. Some birds in the colony could be aggressive, but not him.

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Later in life, Mr. and Mrs. Greedy became an old married couple, always together at their box or outside. It was “very cute,” Kottyan said. Mr. Greedy outlived the median life expectancy of African penguins of 18 years for both males and females.

Penguin Coast is home to the largest colony of African Penguins in North America and the second largest in the world, said Mike Evitts, a spokesperson for the zoo. African Penguins are an endangered species, he said, so every egg hatched and grown into a genetically diverse, healthy adult is important.

That is the legacy of Mr. Greedy, who might not have deserved the name after all.

(Unless we are talking about nesting material and fish. He was greedy about those.)