Limits to reproduction and seed size-number trade-offs that shape forest dominance and future recovery.

The relationships that control seed production in trees are fundamental to understanding the evolution of forest species and their capacity to recover from increasing losses to drought, fire, and harvest. A synthesis of fecundity data from 714 species worldwide allowed us to examine hypotheses that are central to quantifying reproduction, a foundation for assessing fitness in forest trees. Four major findings emerged. First, seed production is not constrained by a strict trade-off between seed size and numbers. Instead, seed numbers vary over ten orders of magnitude, with species that invest in large seeds producing more seeds than expected from the 1:1 trade-off. Second, gymnosperms have lower seed production than angiosperms, potentially due to their extra investments in protective woody cones. Third, nutrient-demanding species, indicated by high foliar phosphorus concentrations, have low seed production. Finally, sensitivity of individual species to soil fertility varies widely, limiting the response of community seed production to fertility gradients. In combination, these findings can inform models of forest response that need to incorporate reproductive potential.

North American tree migration paced by climate in the West, lagging in the East.

Tree fecundity and recruitment have not yet been quantified at scales needed to anticipate biogeographic shifts in response to climate change. By separating their responses, this study shows coherence across species and communities, offering the strongest support to date that migration is in progress with regional limitations on rates. The southeastern continent emerges as a fecundity hotspot, but it is situated south of population centers where high seed production could contribute to poleward population spread. By contrast, seedling success is highest in the West and North, serving to partially offset limited seed production near poleward frontiers. The evidence of fecundity and recruitment control on tree migration can inform conservation planning for the expected long-term disequilibrium between climate and forest distribution.

Continent-wide tree fecundity driven by indirect climate effects.

Indirect climate effects on tree fecundity that come through variation in size and growth (climate-condition interactions) are not currently part of models used to predict future forests. Trends in species abundances predicted from meta-analyses and species distribution models will be misleading if they depend on the conditions of individuals. Here we find from a synthesis of tree species in North America that climate-condition interactions dominate responses through two pathways, i) effects of growth that depend on climate, and ii) effects of climate that depend on tree size. Because tree fecundity first increases and then declines with size, climate change that stimulates growth promotes a shift of small trees to more fecund sizes, but the opposite can be true for large sizes. Change the depresses growth also affects fecundity. We find a biogeographic divide, with these interactions reducing fecundity in the West and increasing it in the East. Continental-scale responses of these forests are thus driven largely by indirect effects, recommending management for climate change that considers multiple demographic rates.

Response of sugar maple to calcium addition to northern hardwood forest.

Watershed budget studies at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, USA, have demonstrated high calcium depletion of soil during the 20th century due, in part, to acid deposition. Over the past 25 years, tree growth (especially for sugar maple) has declined on the experimental watersheds at the HBEF. In October 1999, 0.85 Mg Ca/ha was added to Watershed 1 (W1) at the HBEF in the form of wollastonite (CaSiO3), a treatment that, by summer 2002, had raised the pH in the Oie horizon from 3.8 to 5.0 and, in the Oa horizon, from 3.9 to 4.2. We measured the response of sugar maple to the calcium fertilization treatment on W1. Foliar calcium concentration of canopy sugar maples in W1 increased markedly beginning the second year after treatment, and foliar manganese declined in years four and five. By 2005, the crown condition of sugar maple was much healthier in the treated watershed as compared with the untreated reference watershed (W6). Following high seed production in 2000 and 2002, the density of sugar maple seedlings increased significantly on W1 in comparison with W6 in 2001 and 2003. Survivorship of the 2003 cohort through July 2005 was much higher on W1 (36.6%) than W6 (10.2%). In 2003, sugar maple germinants on W1 were approximately 50% larger than those in reference plots, and foliar chlorophyll concentrations were significantly greater (0.27 g/m2 vs. 0.23 g/m2 leaf area). Foliage and fine-root calcium concentrations were roughly twice as high, and manganese concentrations twice as low in the treated than the reference seedlings in 2003 and 2004. Mycorrhizal colonization of seedlings was also much greater in the treated (22.4% of root length) than the reference sites (4.4%). A similar, though less dramatic, difference was observed for mycorrhizal colonization of mature sugar maples (56% vs. 35%). These results reinforce and extend other regional observations that sugar maple decline in the northeastern United States and southern Canada is caused in part by anthropogenic effects on soil calcium status, but the causal interactions among inorganic nutrition, physiological stress, mycorrhizal colonization, and seedling growth and health remain to be established.