The reproductive strategies of the heterocarpic annual Calendula arvensis (Asteraceae)

Achene polymorphism and various aspects of the reproductive biology of the annual Calendula arvensis L. (Field marigold), were studied to determine the reproductive strategies of the plant. This species normally produces three types of achene: rostrate, cymbiform and annular. Rostrate and cymbiform achenes are larger and heavier than annular achenes, and are adapted to long-range dispersal (by epizoochory and anemochory, respectively). In contrast, annular achenes are smaller in size and weight, and are adapted to short-range dispersal. Achenes germinate over a broad range of temperatures (both in light and in darkness), exhibiting cymbiform achenes the highest germination percentages and annular achenes the lowest under all conditions tested. A fraction of the three types of achenes exhibit dormancy and presumably enter the soil seedbank. Achene types adapted for long-range dispersal (rostrate and cymbiform achenes) produce seedlings that are best able to emerge from deeper burial depths, and that are initially stronger and exhibit earlier flowering than the plants from the annular achenes (which are likely to disperse over shorter distances). These features, together with the fact that fruiting occurs even in the absence of pollinators (automatic geitonogamy), the different mechanisms for achene dispersal (zoochory, anemochory and myrmerochory), and the extended germination, flowering and fruiting periods, facilitate establishment and expansion of this species in unpredictable and disturbed habitats.