Adaptive significance of the prolonged diapause in the western Mediterranean lycaenid butterfly Tomares ballus (Lepidoptera: Lycaenidae)

Diapause is a common dormancy strategy exhibited by many species of invertebrates and insects to temporarily avoid seasonally recurring unfavourable conditions for their development, most usually in winter. Less frequently, a prolonged diapause lasting two or more years is described in species living in unpredictable environments where it is adaptive, but with significant costs. In this paper we examine the occurrence of prolonged diapause in the lycaenid butterfly Tomares ballus. Pupae of this species undergo an obligate diapause from mid-May to late January the following year. However, during our rearing experiments (from 2009 to 2016) the emergence of adults occurred sequentially and a fraction of the pupae remained in diapause for up to seven years. The annual percentage emergence after the first year of diapause was 45.6%, and only barely exceeded 50.0% in 2015. Remarkably, 12 pupae (11.4% of the initial brood) remained in diapause in their eighth year. The negative exponential equation fitted to the emergence data suggests that further emergences may occur within the next five years. Therefore, the potential for successful prolonged diapause of T. ballus pupae may be more than 10 years. The adaptive value of this strategy is discussed in relation to the effects of adverse and unpredictable weather during the flight period of the butterfly, intra-guild competition, parasitoids and changes in habitat quality. We suggest that this strategy may also be exhibited by other species of Mediterranean lycaenids.