Since long before the name "colony collapse disorder" was given to the phenomenon in 2006, scientists have known that honey bee populations in Europe and America were on the decline.
"But we need to distinguish between long-term decline and short-term loss," U.S. Department of Agriculture researcher Bob Danka told a state meeting of beekeepers at Clemson University last month. "Around the world, we are actually gaining bees."
Danka's work in honey bee breeding and genetics is focused on combatting the decline in U.S. hives, which has been precipitous.
"The United States manages only 40 percent of the bees that it did after World War II," he said. "Bee health is declining for various reasons: loss of habitat, pesticides, pests and diseases."
Understanding the dynamics of bee populations and learning to manage them draws members of the South Carolina Beekeepers Association to Clemson twice each year.
"One of the main issues we face is colony collapse disorder. We're losing bees right and left," said Bill Grayson of Laurens County, incoming president of the state association. "We depend on universities like Clemson to help us develop practices that can maintain healthy hives."
Training offered through the meeting included courses in the South Carolina Master Beekeeper program, jointly sponsored by the Clemson Extension Service and the beekeepers association. With four levels of certification, the program broadens beekeepers' knowledge and skills in working with the insects, their hives and the honey they produce.
"Education is important to any industry, and especially to this one," said Danny Howard, a Clemson Extension agent in Greenville County who works with a large number of Upstate beekeepers. "The association pulls together the best information on managing bees so that its members can take it back and put it into practice.
"These are niche marketers; most of them part-time, small businesses that are very important to the state's economy, not only for the honey they produce, but for the pollination they provide," he said. "Honey bees will forage for miles beyond their hives, which makes them crucial to vegetable production, from commercial farms to our home gardens."
Danka points to a host of factors in the United States that converged to put pressure on bee populations. The diverse habitats of a once agrarian culture have given way to cities and subdivisions, leading to declines in bee nutrition.
Even in rural areas, as American agriculture evolved to maintain fewer, larger farms, vast areas of crops that don't depend on bee pollination, such as corn, left less food for honey bees. Pesticides, including those used on home gardens, also have taken a toll.
"We try to educate home gardeners not to dust their gardens. Powdered Sevin dust is an absolute disaster for bees," Grayson said. "Liquid Sevin is much more friendly, especially if gardeners will spray it late in the evening after bees have returned to the hive. By morning it'll be safe when the bees return."
But the single largest contributor may be another insect, a tiny arthropod parasite. Named for an ancient Roman beekeeper, the Varroa mite attaches itself to the honey bee and sucks away its bodily fluid. It also can carry disease that can devastate a hive.
"Varroa is the major driver of over-wintering losses," said USDA's Danka. "Trying to breed bees resistant to varroa mites is like trying to breed sheep resistant to wolves."
Danka and others at Clemson led the beekeepers through ongoing research and management practices to contend with the pest, including the use of Russian bees that appear to have partial resistance to mites. Other tools include encouraging a natural trait of bees to detect and eliminate the mite as they infect bee pupae.
And the beekeepers, an optimistic lot by nature, still find a silver lining even in their adversity: The threats to honey bee populations have made others in agriculture stand up and take notice.
"All my life, beekeeping was the redheaded stepchild of agriculture," said Lawrence Cutts, a lifelong beekeeper and retired chief of the apiary regulatory program at the University of Florida. "Then colony collapse disorder came along and there was a shortage of bees for pollination, and agriculture realized once the bees go away, everything goes away.
"Then we went to the top of the pile when it came to things like getting the legislation and research we need," he said. "Now we are Cinderella at the ball, dancing with the prince."