June Foray, an actress of a thousand voices, who portrayed Rocky the Flying Squirrel and the spy Natasha Fatale on the satirical animated adventures of Rocky and Bullwinkle in the 1960s and other animated creatures and characters on TV and film has died at 99.

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June Foray, an actress of a thousand voices, who portrayed Rocky the Flying Squirrel and the fiendish spy Natasha Fatale on the wickedly satirical animated adventures of Rocky and Bullwinkle in the 1960s and myriad other animated creatures and characters on television and film, died Wednesday in Los Angeles. She was 99.

Her niece Lauren Marems confirmed the death.

Ms. Foray began her remarkable 85-year career by playing an elderly woman in a radio drama in 1929 at age 12. She portrayed scores of radio characters in the 1930s and 1940s. Over the next 60 years, she provided voices for animated shorts, feature films and television shows, as well as record albums, video games, even talking toys. Her last performance was as Rocky in a 2014 Rocky and Bullwinkle cartoon produced by DreamWorks Animation.

Often compared to Mel Blanc, the cartoon virtuoso who supplied the voices of Bugs Bunny, Daffy Duck and Porky Pig, Ms. Foray cackled, chirped, meowed and sometimes sang her way through nearly 300 animated productions, often playing several parts at once with quick shifts of accent, dialect and personality. Her work, unlike that of Blanc, was often uncredited, particularly in her early years.

But her output was prodigious. While she was not well known to the public, the entertainment world called her the First Lady of Animated Voicing. At 94, she became the oldest person to win an Emmy, cited for her Mrs. Cauldron on “The Garfield Show,” and in 2013, she received an Emmy Governors Award.

“June Foray is not the female Mel Blanc,” said Chuck Jones, the legendary animator who proposed her star on Hollywood’s Walk of Fame. “Mel Blanc was the male June Foray.”

On the big screen, she was Lucifer the cat in Walt Disney’s “Cinderella”; a mermaid and a squaw in “Peter Pan” (1953); and Wheezy Weasel and Lena Hyena in “Who Framed Roger Rabbit.” On television, she was Cindy-Lou Who in “How the Grinch Stole Christmas”; Ursula in “George of the Jungle”; and Aunt May Parker in “Spider-Man and His Amazing Friends.”

She also breathed sinister spirit into a doll in a memorable 1963 “Twilight Zone” episode, telling a girl’s stepfather, played by Telly Savalas, “My name is Talky Tina, and I’m going to kill you.”

Ms. Foray portrayed grannies, witches, a fortuneteller, innocent girls, sultry femmes and menageries of anthropomorphic chipmunks, cats, woodpeckers, mice, beagles and other cartoon characters in the adventures of Tom and Jerry, Bugs Bunny, Mr. Magoo, Sylvester and Tweety, Yogi Bear, the Flintstones, the Incredible Hulk, the Smurfs and the Simpsons.

But the cognoscenti said she was at her peak for Rocket J. Squirrel (aka Rocky the Flying Squirrel) and his curvaceous adversary, Natasha Fatale, on the proudly two-dimensional cliffhanger chronicles of “Rocky and His Friends” (later “The Bullwinkle Show”) from 1959 to 1964. In other segments, she played Nell Fenwick, the prim girlfriend of the handsome, muddle-headed Mountie Dudley Do-Right.

Ms. Foray herself said Rocky — with his trademark exclamation “Hokey Smoke!” — was her favorite.

“Everybody asks me that,” she said iin2000. “I think the fans kind of answer that for me. Everybody loves Rocky. I get letters from Belgium, Germany, all over. People don’t think of him as a squirrel. They think of him as a person. And he’s a good little person.”

As the Cold War was heating up and the Red scare turned everyone blue, the gifted voices of Ms. Foray, Bill Scott, Paul Frees and William Conrad gave vivacity to Rocky, a plucky little rodent with an aviator helmet, and his antlered, dimwitted moose pal (Scott) as they battled the inept Slavic schemers Boris Badenov (Frees) and Natasha in Frostbite Falls, Minnesota, a neverland where silliness and puns live forever.

There was plenty of action for the children, slam-bang stories with standard animation gags like characters blowing up or falling out of windows. But on another level, it was satire, parody and rapid-fire wordplay. Dorson Belles warns his radio audience that invaders from outer space are no joke and that everyone should panic. A mystery gas called “votane” turns Democrats into Republicans, and vice versa.

“If you can’t believe what you read in the comic books,” Rocky asks, “what can you believe?”

The Russified Natasha, a villain of many slinky disguises, appears as an Indian princess, Bubbling Spring That Runs in the Meadow. “Call me Bubbles,” she purrs.

No pun was too awful, no malaprop too shameless. Rocky trained at Cedar Yorpantz Flying School. Bullwinkle’s alma mater was Wossamotta U. A jeweled toy boat, the Ruby Yacht of Omar Khayyám, sailed across Veronica Lake. “For a powerful magnate,” Rocky tells a tycoon, “you sure don’t pick up things too quickly.” In one episode, the heroes track a monstrous whale, Maybe Dick.

Besides matching wits with menacing Boris (“Keel Moose!”) and Natasha (“Boris, dollink!”), Rocky and Bullwinkle battle metal-chomping Moon Mice devouring America’s TV antennas. They discover the anti-gravitational element Upsidasium. And the narrator (Conrad) solemnly urges fans to tune in for the next exciting episode: “All in Fever Say Aye, or the Emotion Is Carried,” “The Show Must Go On, or Give ‘em the Acts,” and “Trans-Atlantic Chicken, or Hens Across the Sea.”

After 150 episodes, first on ABC and then on NBC, the series, created by Jay Ward and written by Scott and others, was canceled. But it had a huge cult following. Network reruns aired until 1973 and again in 1981-82. Cable reruns ran through the 1990s. Tributes were held at film festivals. Walt Disney bought videocassette rights for $1 million. The shows were syndicated in the United States, Australia, England and Japan.

PBS produced a documentary, “Of Moose and Men: The Rocky and Bullwinkle Story,” and Ms. Foray provided the voice of Rocky again in 2000 in “The Adventures of Rocky and Bullwinkle,” a feature that combined live action and computer animation. (Rene Russo played Natasha.)

“It seems we’re going to corrupt another generation,” Ms. Foray said at the time.

June Lucille Forer was born in Springfield, Massachusetts, on Sept. 18, 1917, to Maurice and Ida Forer. A high-school speech teacher with a radio program put her on the air.

After her family moved to Los Angeles, she wrote and acted all the parts on her own radio show, “Lady Make Believe,” as a teenager and was soon doing voice-overs for film studios. In the 1940s, she provided voices for a live-action series of film shorts called “Speaking of Animals” and appeared on radio shows starring Danny Thomas, Steve Allen, Jimmy Durante and the team of Phil Harris and Alice Faye.

Her first marriage, to Bernard Barondess in 1941, ended in divorce. In 1955, she married Hobart Donavan, who died in 1976.

Ms. Foray, who lived in the Woodland Hills neighborhood of Los Angeles, leaves no immediate survivors.

In the 1970s, she was president of ASIFA, the international animated film society, which named an award in her honor. She taught voice acting at the University of Southern California in the 1980s, and for decades was a governor of the Academy of Motion Picture Arts and Sciences.