Understanding tree physiological responses to fire is needed to accurately model post‐fire carbon processes and inform management decisions. Given trees can die immediately or at extended time periods after fire, we combined two experiments to assess the short‐ (one‐day) and long‐term (21‐months) fire effects on Pinus ponderosa sapling water transport. Native percentage loss of conductivity (nPLC), vulnerability to cavitation and xylem anatomy were assessed in unburned and burned saplings at lethal and non‐lethal fire intensities. Fire did not cause any impact on nPLC and xylem cell wall structure in either experiment. However, surviving saplings evaluated 21‐months post‐fire were more vulnerable to cavitation. Our anatomical analysis in the long‐term experiment showed that new xylem growth adjacent to fire scars had irregular‐shaped tracheids and many parenchyma cells. Given conduit cell wall deformation was not observed in the long‐term experiment, we suggest that the irregularity of newly grown xylem cells nearby fire wounds may be responsible for decreasing resistance to embolism in burned plants. Our findings suggest that hydraulic failure is not the main short‐term physiological driver of mortality for Pinus ponderosa saplings. However, the decrease in embolism resistance in fire‐wounded saplings could contribute to sapling mortality in the years following fire.

The combination of drought and fire can cause drastic changes in forest composition and structure. Given the predictions of more frequent and severe droughts and forecasted increases in fire size and intensity in the western United States, we assessed the impact of drought and different fire intensities on Pinus ponderosa saplings. In a controlled combustion laboratory, we exposed saplings to surface fires at two different fire intensity levels (quantified via fire radiative energy; units: MJ m−2). The recovery (photosynthesis and bud development) and mortality of saplings were monitored during the first month, and at 200- and 370-days post-fire. All the saplings subjected to high intensity surface fires (1.4 MJ m−2), regardless of the pre-fire water status, died. Seventy percent of pre-fire well-watered saplings recovered after exposure to low intensity surface fire (0.7 MJ m−2). All of the pre-fire drought-stressed saplings died, even at the lower fire intensity. Regardless of the fire intensity and water status, photosynthesis was significantly reduced in all saplings exposed to fire. At 370 days post-fire, burned well-watered saplings that recovered had similar photosynthesis rates as unburned plants. In addition, all plants that recovered or attempted to recover produced new foliage within 35 days following the fire treatments. Our results demonstrate that the pre-fire water status of saplings is an important driver of Pinus ponderosa sapling recovery and mortality after fire.

Summary Wildfires are a global crisis, but current fire models fail to capture vegetation response to changing climate. With drought and elevated temperature increasing the importance of vegetation dynamics to fire behavior, and the advent of next generation models capable of capturing increasingly complex physical processes, we provide a renewed focus on representation of woody vegetation in fire models. Currently, the most advanced representations of fire behavior and biophysical fire effects are found in distinct classes of fine‐scale models and do not capture variation in live fuel (i.e. living plant) properties. We demonstrate that plant water and carbon dynamics, which influence combustion and heat transfer into the plant and often dictate plant survival, provide the mechanistic linkage between fire behavior and effects. Our conceptual framework linking remotely sensed estimates of plant water and carbon to fine‐scale models of fire behavior and effects could be a critical first step toward improving the fidelity of the coarse scale models that are now relied upon for global fire forecasting. This process‐based approach will be essential to capturing the influence of physiological responses to drought and warming on live fuel conditions, strengthening the science needed to guide fire managers in an uncertain future.

Summary The plume of hot gases rising above a wildfire can heat and kill the buds in tree crowns. This can reduce leaf area and rates of photosynthesis, growth, and reproduction, and may ultimately lead to mortality. These effects vary seasonally, but the mechanisms governing this seasonality are not well understood. A trait‐based physical model combining buoyant plume and energy budget theories shows the seasonality of bud necrosis height may originate from temporal variation in climate, fire behaviour, and/or bud functional traits. To assess the relative importance of these drivers, we parameterized the model with time‐series data for air temperature, fireline intensity, and bud traits from Pinus contorta, Picea glauca, and Populus tremuloides. Air temperature, fireline intensity, and bud traits all varied significantly through time, causing significant seasonal variation in predicted necrosis height. Bud traits and fireline intensity explained almost all the variation in necrosis height, with air temperature explaining relatively minor amounts of variation. The seasonality of fire effects on tree crowns appears to originate from seasonal variation in functional traits and fire behaviour. Our approach and results provide needed insight into the physical mechanisms linking environmental variation to plant performance via functional traits.

Summary Disruption of photosynthesis and carbon transport due to damage to the tree crown and stem cambial cells, respectively, can cause tree mortality. It has recently been proposed that fire‐induced dysfunction of xylem plays an important role in tree mortality. Here, we simultaneously tested the impact of a lethal fire dose on nonstructural carbohydrates (NSCs) and xylem hydraulics in Pinus ponderosa saplings. Saplings were burned with a known lethal fire dose. Nonstructural carbohydrates were assessed in needles, main stems, roots and whole plants, and xylem hydraulic conductivity was measured in the main stems up to 29 d postfire. Photosynthesis and whole plant NSCs declined postfire. Additionally, all burned saplings showed 100% phloem/cambium necrosis, and roots of burned saplings had reduced NSCs compared to unburned and defoliated saplings. We further show that, contrary to patterns observed with NSCs, water transport was unchanged by fire and there was no evidence of xylem deformation in saplings that experienced a lethal dose of heat from fire. We conclude that phloem and cambium mortality, and not hydraulic failure, were probably the causes of death in these saplings. These findings advance our understanding of the physiological response to fire‐induced injuries in conifer trees.