NEW YORK, United States — Holiday shoppers spent less money over the Black Friday weekend than in 2015, another sign that US consumers remain wary about opening their wallets without deep discounts.

Shoppers spent an average of $289.19 over the four-day weekend, the National Retail Federation said on Sunday, citing a survey conducted by Prosper Insights & Analytics. Consumers shelled out $299.60 in 2015, the trade group found.

The weekend was characterised by heavy markdowns and a shift toward e-commerce, meaning there were smaller crowds at the mall. Forty-four percent of consumers did their shopping online, compared with 40 percent in brick-and-mortar stores.

“Over one-third of shoppers said 100 percent of their purchases were on sale,” NRF chief executive officer Matthew Shay said in a statement. That increased more than threefold from last year.

By Nick Turner and Jing Cao.