The cause of CCD is currently under investigation. At this point, almost every possible and realistic cause remains a possibility. The following list of leading candidates--in no particular order--is not comprehensive and may change as new information becomes available. Such changes could result in the addition or exclusion of any of the following potential causes.
For more detailed information about each possible cause, see the original publication, Colony Collapse Disorder (CCD) in Honey Bees.
Traditional bee pests and diseases.
Most likely they are not responsible for CCD, as they do not have a history of producing those symptoms, but they may exacerbate the problem.
Bee management. Management style is a broad category, but it can include the type of income pursued with bees--honey production, pollination services--or the routine colony management that beekeepers perform--splitting hives, swarm control, chemical use.
Queen source. Lack of genetic diversity and lineage of bees--both related to queen quality--are being investigated as possible causes of CCD.
Chemical use in bee colonies. Chemicals used to control bee maladies have a variety of sub-lethal effects on all honey bees--workers, queens, and drones--even when the chemicals were used according to label and in accordance with best management practices suggested by specialists.
Chemical toxins in the environment. In addition to being exposed to toxins while foraging, honey bees may also encounter toxins by drinking water contaminated with chemical runoff, encountering various household or commercial chemicals outside of the hive, or via direct inhalation.
Genetically modified crops. Many seeds from which genetically modified crops are grown are dipped first in systemic insecticides that later may appear in the plants' nectar and pollen, making genetically modified plants suspect because of their chemical treatment history, not because of their modified genetics.
Varroa mites and associated pathogens. Varroa mites remain the world's most destructive honey bee killer, so they, the viruses they transmit, and the chemical response they elicit from beekeepers are considered possible causes of CCD.
Nutritional fitness. Malnutrition causes stress to bees, possibly weakening the bees' immune systems. Weak immune systems can affect bees' ability to fight pests and diseases.
Undiscovered or new pests and diseases. Undiscovered, unidentified, or recently-introduced pests and pathogens are considered possible causes of CCD.
Many scientists believe that CCD is caused by some combination of the factors above. For example, some dead bees showing CCD symptoms have had elevated levels of normally-benign pathogens in their bodies, possibly indicating a compromised immune system.
In theory, any stress or combination of stresses--such as chemicals, genetic bottlenecks, or varroa--can suppress a bee's immune system. Considering synergistic effects as a potential cause of CCD makes the disorder increasingly more difficult to study.
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