The Park Grass Experiment at Rothamsted in southeast England was started in 1856, making it the longest-running experiment in plant ecology anywhere in the world. Experimental inputs include a range of fertilizers (nitrogen, phosphorus, potassium, and organic manures) applied annually, with lime applied occasionally, and these have led to an increase in biomass and, where nitrogen was applied in the form of ammonium sulfate, to substantial decreases in soil pH. The number of species per plot varies from three to 44 per 200 m 2 , affording a unique opportunity to study the determinants of plant species richness and to estimate the effect sizes attributable to different factors. The response of species richness to biomass depends on the amount and type of nitrogen applied; richness declined monotonically with increasing biomass on plots receiving no nitrogen or receiving nitrogen in the form of sodium nitrate, but there was no relationship between species richness and biomass on plots acidified by ammonium sulfate application. The response to lime also depended on the type of nitrogen applied; there was no relationship between lime treatment and species richness, except in plots receiving nitrogen in the form of ammonium sulfate, where species richness increased sharply with increasing soil pH. The addition of phosphorus reduced species richness, and application of potassium along with phosphorus reduced species richness further, but the biggest negative effects were when nitrogen and phosphorus were applied together. The analysis demonstrates how multiple factors contribute to the observed diversity patterns and how environmental regulation of species pools can operate at the same spatial and temporal scale as biomass effects.

It is an axiom of life-history theory that reproduction involves age-specific costs in terms of survival or future reproduction. The measurement of costs of reproduction in plants is difficult, and few field studies have measured these costs in terms of fitness or demographic components, thus creating a hiatus between theory and data. In this article, we describe methods for overcoming the problem, illustrated by a field study of balsam fir. We used serial correlation and a permutation test to detect growth costs of reproduction and show how these translate into demographic costs when relative tree size (and therefore growth) is critical to survival. Using chronosequences, we reconstructed the age- and size-specific dynamics of a subalpine population of Abies balsamea. A matrix model describing these dynamics was then used to estimate age- and size-specific probabilities of future survival to maturity ([Formula: see text]). By using a regression model of the relationship between tree size, age, and [Formula: see text], we were able to estimate the maximum age-specific demographic cost of reproduction for trees of all ages. The shape of the age-specific cost curve for A. balsamea may explain why, contrary to a previously published hypothesis, age at first reproduction in A. balsamea does not vary between wave-regenerating and normal populations.

The Park Grass Experiment (PGE) is the longestobserved set of experimental plant communities in existence. Although the gross composition of the vegetation was at equilibrium over the 60-yr period from 1920 to 1979, annual records show that individual species exhibited a range of dynamics. We tested two hypotheses to explain why some species initially increased and why subsequently some of these (the outbreak species) decreased again. The study was designed around eight phylogenetically independent contrasts (PICs), each containing related species with different dynamics. Our first hypothesis was that persistent increasers and outbreakers have higher intrinsic rates of natural increase than control species (species without trends), allowing them to spread when interspecific competition is reduced by drought. This was tested by measuring establishment and seed production of species in field experiments, with and without interspecific competition. Seed production in outbreak species responded more strongly to release from interspecific competition than it did in either of the other groups of species. Our second hypothesis was that outbreak species eventually declined because they lacked the genetic variation necessary to adapt to the novel habitats to which they had initially spread. We tested this by measuring mating systems and genetic diversity in persistent and outbreak species in the PGE. In seven out of seven PICs tested, the outbreak species was more selfing than its persistent relative. There was a significant positive correlation between outcrossing rate and gene diversity. These results support roles for both ecological and genetic traits in long-term dynamics.