Limited evidence for CO 2 -related growth enhancement in northern Rocky Mountain lodgepole pine populations across climate gradients.

Forests sequester large amounts of carbon annually and are integral in buffering against effects of global change. Increasing atmospheric CO2 may enhance photosynthesis and/or decrease stomatal conductance (gs ) thereby enhancing intrinsic water-use efficiency (iWUE), having potential indirect and direct benefits to tree growth. While increasing iWUE has been observed in most trees globally, enhanced growth is not ubiquitous, possibly due to concurrent climatic constraints on growth. To investigate our incomplete understanding of interactions between climate and CO2 and their impacts on tree physiology and growth, we used an environmental gradient approach. We combined dendrochronology with carbon isotope analysis (δ13 C) to assess the covariation of basal area increment (BAI) and iWUE over time in lodgepole pine. Trees were sampled at 18 sites spanning two climatically distinct elevation transects on the lee and windward sides of the Continental Divide, encompassing the majority of lodgepole pine's northern Rocky Mountain elevational range. We analyzed BAI and iWUE from 1950 to 2015, and explored correlations with monthly climate variables. As expected, iWUE increased at all sites. However, concurrent growth trends depended on site climatic water deficit (CWD). Significant growth increases occurred only at the driest sites, where increases in iWUE were strongest, while growth decreases were greatest at sites where CWD has been historically lowest. Late summer drought of the previous year negatively affected growth across sites. These results suggest that increasing iWUE, if strong enough, may indirectly benefit growth at drier sites by effectively extending the growing season via reductions in gs . Strong growth decreases at high elevation windward sites may reflect increasing water stress as a result of decreasing snowpack, which was not offset by greater iWUE. Our results imply that increasing iWUE driven by decreasing gs may benefit tree growth in limited scenarios, having implications for future carbon uptake potential of semiarid ecosystems.