Neurotoxic alkaloid in pollen and nectar excludes generalist bees from foraging at death-camas, Toxicoscordion paniculatum (Melanthiaceae)

Abstract

Abstract Many plants produce broadly active toxins to which specialist herbivores—typically insects—have evolved counter-adaptations, sometimes spawning co-evolutionary arms races. Many non-social bee species are likewise taxonomic host specialists, but the specialists’ pollen hosts frequently attract diverse floral generalists as well, even to flowers of plants that are otherwise chemically defended. In this study of foothills death-camas, Toxicoscordion paniculatum (Nutt.) Rydberg (formerly Zigadenus), we show that its pollen and nectar both contain zygacine, the steroidal alkaloid responsible for this plant’s notorious mammalian toxicity. Hungry naïve adults of a generalist solitary bee, Osmia lignaria Say (Megachilidae), would briefly drink death-camas nectar or biologically relevant doses of zygacine in syrup, followed by prolonged bouts of irritable tongue grooming; many became paralyzed and some even died. Larvae fed dosed provision masses likewise often ceased feeding and sometimes died. Prolonged irritation and subsequent deterrence of foraging O. lignaria likely illustrates why it and 50+ other vernal bee species were absent from death-camas flowers in a five-state survey. The sole visiting bee, Andrena astragali, foraged exclusively at death-camas flowers for pollen and nectar. Thus, a toxic alkaloid found in death-camas pollen and nectar deters generalist bees from flowers of this pollinator-dependent monocot, restricting visitation to a single specialist bee that tolerates death-camas toxins and is its likely pollinator.