• food conditions;
  • litter size;
  • mortality;
  • reproductive success;
  • sex ratio


  • 1
     We studied how differences in the cost of producing male and female offspring in humans affected the productivity of twin vs. singleton deliveries in two ecologically different areas of pre-industrial (1752–1850) Finland. Given the higher energy requirements of male infants, we predicted sons to suffer more from increased litter size and food scarcity than daughters.
  • 2
     We found that the number of offspring surviving to adulthood from a twin delivery differed between the archipelago and mainland areas of rural Finland. On the mainland areas, where crop failures and subsequent famines were common throughout the centuries, twin deliveries were much less successful than in the south-west archipelago, where food conditions were traditionally more stable and survival was ensured by fishing.
  • 3
     Productivity of twin deliveries was modified by the gender composition of the twins; female–female twin births were generally most successful and male–male births least successful.
  • 4
     On the mainland, giving birth to twins of any gender composition never increased the mothers’ reproductive success beyond giving birth to either a male or female singleton, whereas in the archipelago mothers could increase their reproductive output by producing twins. This was because in the archipelago female–female twin deliveries contributed on average more than one adult offspring to the breeding population, whereas a singleton delivery of either gender produced only about 0·7 adults.
  • 5
     Our results show that increases in litter size and variability of food conditions increased male mortality. High female twin survival in the archipelago led to higher fitness of twinning mothers in relation to mothers with only singletons. Twinning has traditionally been significantly more common in the archipelago as compared to the mainland, but contrary to the prediction, there were no apparent differences in the frequency of male–male and female–female twins being born.