Why wait? Trait and habitat correlates of variation in germination speed among Kalahari annuals

The longer soil has to stay moist to allow germination the more likely that seedlings experience favourable moisture conditions. Since theory predicts that fitness variance-reducing traits will be negatively correlated, we tested the hypothesis that time to germination is negatively correlated with the ability of radicle growth to keep up with the drying front. We measured time to germination and root elongation rate (RER) in 14 Kalahari annuals. We controlled for habitat (canopy association and sand content), germinability, median base water potential for germination (psi(50)), seed mass and seed shape as a persistence surrogate. For species and phylogenetically independent contrasts (PICs), we did not find a relationship between time to germination and RER. However, we found a negative relationship of time to germination with RER for PICs when controlling for sand content and psi(50). Seed shape increased with time to germination which can be explained by reduced opportunities for germination in slow-germinating species that select for persistence. We found a positive relationship between time to germination and psi(50), suggesting a continuum of risky to cautious germination. psi(50) was not correlated with RER suggesting that variation in psi(50) reflects different drought-adapted traits. Probably the relationship of time to germination with RER is not mediated by seed mass, which was not correlated with time to germination as found by others, though RER was positively correlated with seed mass. Instead of the seed size-seed number trade-off, a trade-off with resource capture may explain variation in RER: more root hairs or rootlets may increase resource capturing surface while reducing RER. For habitat, we found a (positive) relationship with time to germination only for canopy association. This may be explained by maximization of resource capture at the expense of RER being favoured by the higher nutrient and water availability under canopies. Future studies should clarify which trade-offs govern variation in time to germination, focussing on a possible resource capture-RER trade-off.