Physiological time is important for understanding the development and seasonal timing of ectothermic animals, but has largely been applied to developmental processes that occur during spring and summer such as morphogenesis. There is a substantial knowledge gap in the relationship between temperature and development during winter, a season that is increasingly impacted by climate change. Most temperate insects overwinter in diapause, a developmental process with little obvious morphological change. We used principles from the physiological time literature to measure and model the thermal sensitivity of diapause development rate in the apple maggot fly Rhagoletis pomonella, a univoltine fly whose diapause duration varies substantially within and among populations. We showed that diapause duration could be modelled with simple linear relationships between diapause development rate and temperature. Low temperatures were necessary for most individuals to complete diapause, similar to many insects. However, we also found evidence for an ontogenetic shift in the thermal sensitivity of diapause: diapause development proceeded more quickly at high temperatures later in diapause and in the absence of quiescence, a phenomenon not previously reported. Increasingly warmer temperatures during and after winter may impact the phenology of this and other insects with temperature-sensitive diapause, impacting their viability.
Low temperatures associated with winter can limit the survival of organisms, especially ectotherms whose body temperature is similar to their environment. Important adaptations for overwintering such as cold hardiness and diapause have been well-explored in many insect taxa. However, there is a gap in understanding how overwintering may vary among groups of species that interact closely, such as multiple parasitoid species that attack the same host insect. Our study investigated cold tolerance and diapause phenotypes in three endoparasitoid wasps of the apple maggot fly Rhagoletis pomonella (Diptera: Tephritidae): Utetes canaliculatus, Diachasma alloeum, and Diachasmimorpha mellea (Hymenoptera: Braconidae). Using a combination of respirometry and eclosion tracking, we detected diapause phenotypes in all three wasp species, remarkably similar to the fly host. Weak diapause was rare (< 5%) in all three wasp species, and while most D. mellea (93%) entered prolonged diapause under warm conditions, the majority of U. canaliculatus (92%) and D. alloeum (72%) averted diapause (non-diapause). There was limited interspecific variation in acute cold tolerance among the three wasp species: wasps and flies had similarly high survival (>87%) following exposure to extreme low temperatures (-20 degrees Celsius) as long as their body fluids did not freeze. The wasp species showed little interspecific variation in survival following prolonged exposure to mild chilling of 8 or more weeks at 4 degrees Celsius. This study shows remarkable conservation of cold tolerance and diapause phenotypes within and across trophic levels. The interaction between diapause phenotype and cold hardiness in these parasitoids is an interesting direction for future research.
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