of the common polygynous breeding systems in which big males compete fiercely with each other to acquire a harem of females. Instead, big females compete fiercely to acquire a harem of smaller males, for each of which in turn the female lays a clutch of eggs, and each of which proceeds to do most or all of the work of incubating the eggs and rearing the young. The best known of these female sultans are the shore birds called jacanas (alias lily-trotters), Spotted Sandpipers, and Wilson's Phalaropes. For instance, flocks of up to ten female phalaropes may pursue a male for miles. The victorious female then stands guard over her prize to ensure that only she gets to have sex with him, and that he becomes one of the males rearing her chicks.
Clearly, sex-role-reversal polyandry represents for the successful female the fulfillment of an evolutionary dream. She wins the battle of the sexes by passing on her genes to far more clutches of young than she could rear, alone or with one male's help. She can utilize nearly her full egg-laying potential, limited only by her ability to defeat other females in the quest for males willing to take over parental care. But how did this strategy evolve? Why did males of some shorebird species end up seemingly defeated in the battle of the sexes, as polyandrous co-“husbands,” when males of almost all other bird species avoided that fate or even reversed it to become polygynists?
The explanation depends on shorebirds' unusual reproductive biology. They lay only four eggs at a time, and the young are precocial, meaning that they hatch already covered with down, with their eyes open, and able to run and find food for themselves. The parent doesn't have to feed the chicks but only has to protect them and keep them warm. That's something a single parent can handle, whereas it takes two parents to feed the young of most other bird species.
But a chick that can run around as soon as it hatches has undergone more development inside the egg than the usual helpless chick. That requires an exceptionally large egg. (Take a look sometime at a pigeon's typically small eggs, which produce the usual helpless chicks, to understand why egg farmers prefer to rear chickens with big eggs and precocial chicks.) In Spotted Sandpipers, each egg weighs fully one-fifth as much as its mother; the whole four-egg clutch weighs an astonishing 80 percent of her weight. Although even monogamous shorebird females have evolved to be slightly larger than their mates, the effort of producing those huge eggs is still exhausting. That maternal effort gives the male both a short-term and a long-term advantage if he takes over the not too onerous responsibility of rearing the precocial chicks alone, thereby leaving his mate free to fatten herself up again.
His short-term advantage is that his mate thereby becomes capable of producing another clutch of eggs for him quickly, in case the first clutch is destroyed by a predator. That's a big advantage, because shorebirds nest on the ground and suffer horrendous losses of eggs and chicks. For example, in 1975 a single mink destroyed every nest in a population of Spotted Sandpipers that the ornithologist Lewis Oring was studying in Minnesota. A study of jacanas in Panama found that forty-four out of fifty-two nests failed.
Sparing his mate may also bring the male a long-term advantage. If she does not become exhausted in one breeding season, she is more likely to survive to the next season, when he can mate with her again. Like human couples, experienced bird couples that have worked out a harmonious relationship are more successful at raising young than are bird newly weds.
But generosity in anticipation of later repayment carries a risk, for male shorebirds as for humans. Once the male assumes sole parental responsibility, the road is clear for his mate to uso her free time in whatever way she chooses. Perhaps she'll choose to reciprocate and remain available to her mate, on the