Where winters are severe, aphids reproduce parthenogenetically and viviparously in summer, switch to sexual reproduction in late summer, and produce winter-hardy eggs by the end of the season. The role of day length and temperature in initiating seasonal changes from parthenogenetic to sexual reproduction by pea aphids, Acyrthosiphon pisum (Harris) (Hemiptera: Aphididae), are described and the selection pressures that affect the timing of this transition are investigated. Over four seasons, a pea aphid clone was sampled from field cages through late summer in southern Manitoba, Canada, and reared in the laboratory to determine the phenotypes of progeny produced as the season progressed. The timing of transitions from one phenotype to another under natural day length and temperature, and the critical day lengths that caused the transitions, coincided with expectations from laboratory studies of photoperiodic responses. Males and mating females appeared later when the weather in August was warm than when it was cool. The timing of seasonal changes was adapted to minimize the physiological time to the end of the season, which maximized the number of asexual summer generations. Ambient temperature modulated the response to day length and fine-tuned the timing of sexual reproduction to adapt for annual variation in autumn weather.