SAVANNAH, Ga. — Rare sea turtles nesting on beaches in Georgia and the Carolinas laid eggs at a slower pace this summer after a record-smashing 2016 season.

The nesting season for loggerhead sea turtles that ran from May through August still yielded nest counts well above average, said state biologists who had little hope the turtles would match last year’s impressive performance. They said the latest season gives them more encouragement that the federally protected species may be rebounding.

“We’re still significantly above average,” said Mark Dodd, the biologist who heads the sea turtle recovery program for the Georgia Department of Natural Resources. “We generally have a down year after a couple of big years. And if this is our down year, next year should be really good.”

Volunteers scouring Georgia beaches at daybreak throughout the summer counted 2,141 total loggerhead nests. That’s quite a shortfall compared to the 3,289 nests recorded the previous year.

But sea turtles had a nesting boom in the beaches of the Southeast last year, shattering previous records from North Carolina to Florida. Georgia alone saw a staggering 40 percent jump in total loggerhead nests in 2016. The smaller number of nests seen this summer is still 60 percent higher than the state average in Georgia since 1989.

Neighboring states posted similar numbers.

The South Carolina Department of Natural Resources reported 5,169 loggerhead nests this summer. That’s down from 6,446 nests a year ago — another state record — but still 75 percent greater than South Carolina’s 20-year average. There were 1,181 loggerhead nests tallied over the summer in North Carolina, which hit a record 1,580 nests a year ago.

Loggerhead sea turtles, which can grow up to 300 pounds, are protected by federal law as a threatened species. Each summer, adult females crawl from the surf of the Atlantic Ocean onto beaches to dig nests for their pingpong-ball sized eggs.

During the nesting season, volunteers from North Carolina to Florida comb the shoreline each day around sunrise to catalog new nests and cover them with protective screens to keep out wild hogs and other predators until the eggs hatch.

Nest numbers can fluctuate wildly year-to-year. Biologists say that’s because female loggerheads tend to lay eggs only every three to four years. But experts say counts have improved significantly over the past three decades, likely thanks to conservation efforts such as requiring shrimp boats to use nets equipped with built-in escape hatches for turtles.

Last year, females loggerheads must have had an abundance of food and other perfect conditions that led to the nesting boom of 2016, said Charlotte Hope, a South Carolina state biologist who studies sea turtles.

“I don’t expect to see that just continually go up,” Hope said. “It’s up and down, up and down. … But obviously the trend is going up, instead of down like it was in the 1980s.”

By far the busiest U.S. state for sea turtle nesting, Florida doesn’t keep a running count because of overwhelming numbers. Last year, a record 122,707 loggerhead nests were counted on more than 200 Florida beaches.

Simona Ceriani, a research scientist who studies sea turtles for the Florida Fish and Wildlife Commission, said she also expects lowers numbers in Florida, which doesn’t report numbers until the end of November.