Population ecology on an environmental gradient: Cakile edentula on a sand dune

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Abstract
A naturally-occurring sand dune population of the annual plant Cakile edentula (Brassicaceae) was studied for two years. The plants grew along an environmental gradient stretching from open sand beach (seaward) to densely vegetated dunes (landward). Survivorship and reproductive output were estimated from plants in permanent quadrats. The dispersal of seeds was documented by sifting fruits from the sand substrate at different seasons. Seedlings germinate in April and May and begin flowering in July and August. They may continue to flower until October unless destroyed by autumn winds or heavy frost. Although seaward plants germinate approximately a month later than landward plants, they grow more rapidly and by September may be two orders of magnitude larger than landward plants (dry weight of vegetative parts 6.86±3.97 g compared to 0.029±0.006 g; dry weight of fruits 5.92±4.27 g compared to 0.016±0.005 g; mean with 95% CI). In both years, seedling survivorship and mature plant reproductive output declined significantly with distance landward. The large plants at the seaward end of the gradient produced most of the fruits (144 and 278 fruits per capita in 1975 and 1976 respectively) but a large proportion of these were moved landward by wind and waves during the winter. Thus, at the seaward end of the gradient, the main influx on individuals was from reproduction, and the main loss of individuals was from dispersal landward during the winter. The small plants at the landward end of the gradient produced few fruits (1.8 and 1.2 fruits per capita in 1975 and 1976 respectively), and mortality greatly exceeded this reproductive output. Thus, at the landward end of the gradient, the main loss of individuals was through seedling mortality, but this was balanced by a large annual influx of individuals from the seaward end of the gradient. Plants at the landward end of the gradient therefore exist only because of annual dispersal of seeds landward. Most seeds produced at the seaward end of the gradient disperse from an area of good habitat (high survivorship and high reproductive output) to an area of poor habitat (low survivorship and low reproductive output).