Ethel Louise was the sister with the outgoing, strong personality, while her twin, Samantha, was the follower.

“;They were the tightest sisters you could imagine,”; said Richard Ball, mammal curator at the Honolulu Zoo.

The two African lion sisters, who were born the same day, died together yesterday at the ripe old age of 22.

They were euthanized at the Honolulu Zoo due to age-related health problems including arthritis, cystic livers and nodules in the lungs, said Honolulu Zoo Director Stephen Walker.

The pair were a prime exhibit attraction at the Honolulu Zoo for the past 111/2 years. Until two weeks ago, they were the only lions on exhibit since the 2007 death of 21-year-old male lion, Apollo.

The sisters, who would have turned 23 April 3, lived to the outer range of life expectancy of captive African lions, Walker said, much longer than those in the wild.

“;They're such an iconic species, especially when Apollo was alive,”; general curator Peter Luscomb said. “;He was their big male. People loved the exhibit.”;

Ball said yesterday, “;There were a lot of tears this morning,”; particularly from their keepers.

Signs were posted for zoo-goers explaining what had happened.

The lions were born at the Wild Wilderness Zoo in Arkansas. The Honolulu Zoo bought them from the Pittsburgh Zoo and they moved to Honolulu July 28, 1997.

Ethel Louise was the leader of the two, Ball said. “;Samantha was her tight buddy.”;

“;On evenings when Ethel decided she wouldn't come in, Samantha would stay out with her, and that was to the very end,”; Ball said.

Apollo favored Samantha over Ethel Louise, who wasn't afraid of him.

“;When it came right down to it, Ethel Louise and Samantha were too tight for even a male lion to get between,”; Ball said.

The zoo will be getting a 2-year-old male lion from the San Diego Wild Animal Park at the end of March, and is trying to acquire at least two females.