The Ecology of Seed Heteromorphism in Heterosperma Pinnatum in Central Mexico

Abstract

Heterosperman pinnatum is an annual composite that produces heteromorphic achenes, with the proportion of achene morphs varying geographically. Field, greenhouse, and laboratory experiments were used to examine the ecological consequences of the heteromorphism. Achenes were classified into three types based on morphology. Central, intermediate, and peripheral morphs, named for the positions occupied within the fruiting head, range in shape from long, thin—beaked achene with barbed awns (central) to short wide achenes without beaks or awns (peripheral). Considerable within— and between—population phenotypic variation in achene and head characteristics was documented. Field experiments using artificial dispersal agents showed that central achenes are the most likely to adhere to animals and that the proportion of a population's achenes presented for dispersal that adhere depends on the proportion of central achenes produced in the population and on the proportion of achenes with adhesive awns. Once achenes had reached the ground, mean and median secondary dispersal distances were short (@<20 cm). Central achenes lose innate dormancy earlier than peripheral achenes during the period between autumn achene production and the onset of the summer rainy season. Germination is inhibited by darkness, though darkness becomes less inhibitive at the onset of the germination season (earlier for central achenes). In the laboratory, peripheral achenes germinated over a slightly narrower range of temperatures than central achenes. Percent germination was greater for central than intermediate achenes, and for intermediate than peripheral achenes, in two natural germination events in the field. Seed bank samples indicated that few if any achenes remain dormant between years. No significant differences in the growth rates of young seedlings were found in the greenhouse. A demographic experiment documented trade—offs resulting from the dormancy differences: central achenes tended to germinate earlier than intermediate and peripheral ones, and early germination resulted in greater mortality, but survivors that had germinated earlier attained greater size and produced more seeds. Intermediate achenes, while intermediate in all components of fitness, had the highest total fitness. This was because they resembled peripheral achenes in germinating late and having high survival, but they resembled central achenes in having higher percent germination. The results of these experiments suggest that a achene behavior ranges from "low—risk" (peripheral achenes) to "high—risk" (central achenes) within the progeny of a single individual, while individuals and populations vary in the proportion of offspring exhibiting each type of behavior.