A colorful quartet of black-and-whites

The new DreamWorks animated feature “Madagascar” is filled with eminently watchable creatures. A cuddly lion, frisky zebra, maternal hippo and hypochondriac giraffe (voiced by Ben Stiller, Chris Rock, Jada Pinkett Smith and David Schwimmer, respectively) must fend for themselves for the first time when they escape from the Central Park Zoo and end up on the island of Madagascar.

What gets them there, and sets the fish-out-of-water story in motion, is a fiendish little band (emphasis on little) of characters that have their hearts set on making it to Antarctica. In other words, when it comes time to twist the plot, keep an eye on the penguins.

It wasn’t always thus. When the project was in the early stages, the penguins didn’t even have speaking roles. The route that took them from extras to scene stealers is circuitous -- one might even call it a long and winding road, because the Beatles figure into the story.

Once upon a time, back in 1999, before “Madagascar” was even a gleam in Jeffrey Katzenberg’s eye, the DreamWorks Animation CEO had the concept of a “Hard Day’s Night”-style movie starring penguins. Producer Mireille Soria and director Eric Darnell (“Antz”) began developing this “rockumentary project” about four penguins that made great music together and the effects of fame and success on the group. The filmmakers made a minute-long clip, using an early Beatles press conference for dialogue and creating a set of truly endearing penguins as band members.

Coincidentally, filmmaker Tom McGrath was working on an animated penguin movie of his own, for a small independent production company. His project was a more naturalistic story, about the real world of the birds.

“Penguins are such neat animals, and they’re natural comedians. Just working on that gave me a soft spot in my heart for them,” he said. When that film fell through, after a year and a half of work, McGrath had to move on.

He went to DreamWorks, met with Katzenberg and Soria, and was shown snippets of projects that various crews were working on.

“The last thing that showed up was [a clip of] these penguins talking like Beatles. And I just went, ‘What’s this?’ ” McGrath said. He then met with Darnell, and the two hit it off. “I really wanted to work on the project, and I was excited to come to DreamWorks, so I signed a deal,” McGrath explained.

The week he arrived at DreamWorks, though, the project that first attracted him came to a screeching halt. Though DreamWorks had managed to get the music clearances from three-fourths of the Beatles, George Harrison (and later his estate) balked.

“It kind of broke our hearts when it didn’t come through,” said Darnell.

“I was like, ‘Sigh, no penguins,’ ” McGrath added.

A few months later, Darnell and Soria asked McGrath to be the storyboard artist for “Madagascar.” His job was to sketch visual descriptions of every scene that could then be viewed in sequential order, like a movie, to see how the story played out. “We do this for years, until we feel like we have the best film we can make,” McGrath said.

During this process, one scene was coming across a bit too predictably. When the animals are stored in crates on a ship bound for Africa, a storm comes up, they’re tossed overboard, and they wind up on the beach in Madagascar. McGrath was asked to come up with a less cliched twist than the old storm at sea. His creative process went like this: “If all these animals are going back to Africa, what if some animals didn’t belong there, or even want to go, and it was a clerical error on someone’s part?”

McGrath looked at the penguins at the zoo and thought, “Well, if they were on the boat, they wouldn’t want to go to Africa at all. They’d want to go to Antarctica.” From there he came up with a band of penguins with a POW sensibility, capable of taking over the ship and turning it around.

The scene was such a hit, with the militaristic penguins played against their cute, cuddly type, that Darnell and Soria wanted to use them elsewhere.

And the tuxedoed foursome have been such a hit at DreamWorks that work began on a short film about them, even before “Madagascar” was completed. It is due out this year.

The penguins weren’t the only ones upgraded to larger roles. McGrath went from story artist to co-director as a result of his bright ideas. To Darnell, the promotion was just common sense.

“Frankly, he was doing [the job] before he got it -- there’s a lot of amazing stuff that goes on in that head,” Darnell said. “We wanted to take advantage of it.” (Darnell and McGrath are also the film’s writers, with Mark Burton and Billy Frolick.)

As if that weren’t enough, the penguins also gave McGrath a role in the film. He had been voicing the lead penguin, the tough-as-nails Skipper, in the temporary tracks. Everyone liked his voice so much that when the time came to cast the role, he won it hands down.

The penguins lured other colleagues into the action too. Skipper’s right-hand bird, Kowalski, is played by Chris Miller (Magic Mirror in “Shreks” 1 and 2 and director of “Shrek 3”). Christopher Knights, an assistant editor, plays the eager, lowly Private. And clever little Rico is voiced -- or, rather, expressed -- by an uncredited Katzenberg.

“The irony for us is that he’s the one who doesn’t talk,” said Soria of Rico. “There’s something very Dadaistic about that, isn’t there?”