Sperm competition: evolutionary causes and consequences.

The interaction between functional and mechanistic approaches to sperm competition and between male and female perspectives are described and illustrated by a study of the zebra finch, Taeniopygia guttata. Sperm competition experiments in the laboratory show that last male sperm precedence occurs (as it does in many other taxa) although the mechanism is unknown (as in most other taxa). Empirically-derived values were used to construct a mathematical model of sperm competition in the zebra finch. The model indicates that precedence occurs as a consequence of: (i) the temporal pattern of pair copulations; (ii) the rate at which sperm are lost from the female tract; and (iii) more sperm being transferred during extra-pair copulations than during pair copulations. The latter effect is a consequence of males seeking extra-pair copulations after their own pair copulation period has ended. The effect of sperm numbers on the pattern of sperm precedence may be further increased by: (i) extra-pair males increasing ejaculate size (sperm numbers) (for which there is no evidence); (ii) extra-pair males being of a better quality and transferring more sperm or better quality sperm (for which there is some evidence); and (iii) cryptic female choice. Females eject over 99% of sperm following insemination; if they eject fewer sperm from males chosen as extra-pair copulation partners, the potential for cryptic female choice is considerable. However, this is still being investigated. The model also predicts the optimal time for an extra-pair copulation to occur (from either a male or female perspective). A comparison between the predicted and observed pattern suggests that the optimal timing of extra-pair copulations is constrained in both sexes.