Which of the following statements is true about Fleischer Studios?

Fleischer Studios is well-known for being operated by brothers Max and Dave Fleischer, who played a pivotal role in the development of animation during the early 20th century. The studio was responsible for popular characters and series, including Betty Boop and Popeye, and it contributed significantly to the animation industry through innovative techniques and storytelling. The collaborative dynamic between the brothers was crucial to the studio's creative output and success in an era dominated by other major studios. This familial connection helped shape the studio's identity and influence the direction of animated filmmaking at that time.

The other statements do not accurately reflect the studio's history: it was actually established in New York City rather than California, while some of its animations contained darker themes, it was not primarily known for being dark-themed—there was a range of content produced. Lastly, Fleischer Studios did not strictly produce feature films. Instead, they were known for both short films and series that complemented their features.