Rainfall and the interaction of microclimate with larval resources in the population dynamics of checkerspot butterflies (Euphydryas editha) inhabiting serpentine grassland

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Abstract
The interaction of host plant phenology and microclimatic heterogeneity was examined to determine its role in the population dynamics of checkerspot butterflies, Euphydryas editha, inhabiting serpentine grassland in California's outer Coast Range. Within the 2–3 hectares inhabited by a population of E. editha (Jasper Ridge Area H), microclimatic differences resulting from topographic heterogeneity largely determine the temporal and spatial pattern of senescence of the larval host plants, Plantago erecta and Orthocarpus densiflorus. Survival of larvae from hatching to diapause is extremely low as a result of unpredictable variation in the timing of larval development relative to the timing of host plant senescence, both of which are mediated by microclimatic patterns. During this study, population H declined to near extinction as a result of two consecutive years of record rainfall that apparently disrupted the tenuous temporal relationship between larval development and plant senescence. Retarded development of post-diapause larvae led to a late and extended flight season and delayed egg production; this in turn resulted in massive mortality of pre-diapause larvae due to starvation because host plant senescence occurred before larvae became large enough to enter diapause. Adult population size the following spring was the smallest in 25 years of study. This work emphasizes the importance of microclimatic heterogeneity for understanding population-level processes in small ectothermic animals and underlines the potential importance of such heterogeneity in the establishment of reserves designed to protect such animals