The Financial Industry Regulatory Authority fined five broker-dealers a total of $385,000 for the illegal sale of more than eight billion shares of penny stocks.

Most of those sales involved one penny-stock company, Universal Express Inc. The five broker-dealers sold more than 7.5 billion shares of Univeral’s unregistered stock for proceeds of about $8.4 million.

FINRA announced Tuesday that the firms also failed to take appropriate steps to determine whether the securities could be sold without violating federal registration requirements.

The five firms are New York-based Fagenson & Co. Inc., which reported earning $44,000 in commissions from the sale of unregistered Universal stock (it was fined $165,000); RBC Capital Markets Corp., which earned $68,000 in commissions (fined $135,000); Alpine Securities Corp. of Salt Lake City, which earned $47,000 in commissions (fined $40,000); Boca Raton, Fla.-based Equity Station Inc., which earned $13,575 in commissions (fined $25,000); and Olympus Securities, of Montville, N.J., which earned $5,200 in commissions (fined $20,000).

Alpine and Olympus declined to comment on the fine. The other three did not return phone calls seeking comment.

“Brokerage firms are the first line of defense when it comes to preventing the illegal distribution of unregistered securities into the public markets,” James S. Shorris, and executive vice president and executive director of enforcement at FINRA, said in a press release. “The failure to detect and prevent these sales creates serious risks to the unsuspecting customers who purchased these unregistered securities.”

In settling these matters, the firms neither admitted nor denied the charges, according to FINRA.