Are trade-offs in plant resprouting manifested in community seed banks?

Trade-offs in allocation to resprouting vs. seedling regeneration in plants are predicted to occur along fire disturbance gradients. Increased resprouting ability should be generally favored in plant communities with a high probability of crown fire return. Hence, communities dominated by resprouters are predicted to have smaller seed banks than those dominated by species killed by fire. We tested whether there were trait shifts in resprouting ability among growth forms (short-lived herbaceous vs. ground-dwelling perennials vs. shrubs) and among communities (rocky outcrop vs. sclerophyll forest) with contrasting probabilities of crown fire return. Resprouting was more common in ground-dwelling perennials and in the sclerophyll forest community with a high probability of crown fire. Soil seed banks were sampled in rocky outcrop and sclerophyll forest communities in recently burned (18 months) and long-since-burned (12 years) locations at interspersed replicated sites. Collected seed banks were treated with orthogonal treatments of fire stimuli or no stimuli, and seedling emergence was measured in controlled conditions. Seed bank composition reflected the pattern of extant vegetation, with resprouting species being more common in the community with a higher probability of crown fire. Overall, however, resprouting species were poorly represented in the seed bank compared to those species killed by fire. Predicted shifts in allocation to seed production were strongly manifested in community seed banks across the disturbance gradient. Fewer species, seedlings, and seedlings per adult emerged from seed banks in the sclerophyll forest. This suggests that the dominance of resprouting species influences recruitment at the community scale. Community patterns in the seed bank also reflected predicted trade-offs with plant size and growth rate. Short-lived species that are killed by fire dominated the seed bank on rocky outcrops, while longer-lived resprouting species were found in low abundance. Life history trade-offs in persistence and regeneration strongly contribute to coexistence patterns between and within communities with contrasting probabilities of fire return.     

Growth rates, seed size, and physiology: do small-seeded species really grow faster?

Relative growth rate (RGR) is currently the most commonly used method for measuring and comparing species' intrinsic growth potential. Comparative studies have, for example, revealed that small-seeded species have higher RGR, leading to the common belief that small-seeded species possess physiological adaptations for rapid growth that would allow them to outgrow large-seeded species, given sufficient time. We show that, because RGR declines as individual plants grow, it is heavily biased by initial size and does not measure the size-corrected growth potential that determines the outcome of competition in the long-term. We develop a daily growth model that includes a simple mechanistic representation of aboveground and belowground growth and its dependency on plant size and environmental factors. Intrinsic growth potential is encapsulated by the size-independent growth coefficient, G. We parameterized the model using repeated-harvest data from 1724 plants of nine species growing in contrasting nutrient and temperature regimes. Using information-theoretic criteria, we found evidence for interspecific differences in only three of nine model parameters: G, aboveground allocation, and frost damage. With other parameters shared between species, the model accurately reproduced above- and belowground biomass trajectories for all nine species in each set of environmental conditions. In contrast to conventional wisdom, the relationship between G and seed size was positive, despite a strong negative correlation between seed size and average RGR, meaning that large-seeded rather than small-seeded species have higher size-corrected growth potential. Further, we found a significant positive correlation between G and frost damage that, according to simulations, causes rank reversals in final biomass under daily temperature changes of +/- 5 degrees C. We recommend the wider use of this new kind of plant growth analysis as a better way of understanding underlying differences in species' physiology; but we recognize that RGR is still a useful metric if considering the potential rate of population increase in empty habitats.     

Ecological and evolutionary mechanisms for low seed: ovule ratios: need for a pluralistic approach?

Central to the ecology and evolution of a broad range of plants is understanding why they routinely have submaximal reproduction manifested as low seed : ovule and fruit : flower ratios. We know much less about the processes responsible for low seed : ovule ratios than we do for fruit : flower ratios. Current hypotheses for low seed : ovule ratios are largely drawn from those for fruit : flower ratios, including proximate (ecological) causes of pollen limitation, resource limitation, and pollen quality, as well as the ultimate (evolutionary) hypothesis of "bet hedging" on stochastic pollination. Yet, such mechanisms operating on fruit : flower ratios at the whole-plant level may not best explain low seed : ovule ratios at the individual-flower level. We tested each of these proximate and ultimate causes for low seed : ovule ratios using the specialized pollination mutualism between senita cacti (Pachycereus schottii) and senita moths (Upiga virescens). Seed : ovule ratios were consistently low (approximately 0.61). Such excess ovule production by senita likely has a strong genetic component given the significant differences among plants in ovule number and the consistency in ovule production by plants within and among flowering seasons. Excess ovule production and low seed : ovule ratios could not be explained by pollen limitation, resource limitation, pollen quality, or bet hedging. Nevertheless, phenotypic selection analyses did show significant selection gradients for increased ovule number, suggesting that other evolutionary processes may be responsible for excess ovule production and low seed : ovule ratios. In contrast, low fruit : flower ratios at the whole-plant level were explained by an apparent equilibrium between pollen and resource limitation. Thus, mechanisms responsible for low fruit : flower ratios at the whole-plant level are not necessarily in accord with those of low seed : ovule ratios at the individual-flower level. This suggests that we may need to adopt a more pluralistic approach to seed : ovule ratios and consider alternative hypotheses, including a greater array of proximate and ultimate causes. Initial results of this study suggest that floral allometry, selection on correlated floral traits, stigma clogging with pollen grains, and style clogging with pollen tubes may provide promising avenues for understanding low seed : ovule ratios.