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Queens do “age,” and in the process may exhibit poor brood patterns, suggesting that they be replaced. In general, few queens are good for more than a year and a half of egg‐laying, by which time their spermatheca simply runs out of viable sperm. At that point some eggs laid in worker cells will develop into drones (Figure 5.27).

The workers apparently cue on unfertilized eggs as a signal that the queen is failing and will usually supersede a queen before this becomes noticeable, but if they wait too long, they may not have any female larvae from which to raise a replacement queen.

Figure 5.26 A nice egg‐laying pattern by a queen – each egg centered in its cell, with no misses. The eggs are glued upright and typically slightly tilted (often the eggs are all tilted in the same direction). A pattern such as this indicates a high‐quality queen.

Figure 5.27 Raised drone cells scattered among worker cells, as opposed to groups of drone cells at the edge of the comb, indicate that a queen has gone “drone layer” – meaning that she is running out of stored sperm with which to fertilize worker eggs.