Differences in branch hydraulic architecture related to the aridity of growing sites and seed sources of coastal Douglas-fir saplings.

To better understand hydraulic adaptations of coastal Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) to local climate, we examined genetic (G) and environmental (E) responses of branch hydraulic architecture of 7-year-old saplings from dry and wet climates of origin grown at a relatively dry and a relatively wet common garden site in western Oregon. We sampled 2 years of branch growth from three dry-source and three wet-source families grown at both sites (72 branches, total). Overall, only 4 of the 11 traits had significant genetic (G) effects, whereas 9 traits had significant environmental (E) effects (P < 0.05). Both dry and wet sources had higher leaf-specific conductance (kl) at the dry than the wet site, but the values were achieved by different mechanisms and driven by G × E effects for leaf area/sapwood area (Al/As), shoot length (L), specific conductivity (Ks) and leaf-specific conductivity (Kl). Dry sources achieved higher kl in the dry site through higher Kl (via a lower Al/As and no change in Ks) with no difference in L. Wet sources achieved higher kl at the dry site through no difference in Kl (via no effect on Al/As, despite decreases in Al and As, and lower Ks) with lower L. Vulnerability to embolism (measured as percentage loss of conductivity at 4 MPa) had no G effect but an E effect, with slightly lower values at the dry site. Specific leaf area had G and E effects, with lower values for the dry sources and site. There were no G or E effects on wood density. The different responses of dry and wet sources to site aridity suggest that populations are differentially adapted to the aridity of growing sites. Population variation in response to aridity should be considered when selecting seed sources for establishing forests for future climates.

Projected impacts of climate change on the range and phenology of three culturally-important shrub species.

Climate change is shifting both the habitat suitability and the timing of critical biological events, such as flowering and fruiting, for plant species across the globe. Here, we ask how both the distribution and phenology of three food-producing shrubs native to northwestern North America might shift as the climate changes. To address this question, we compared gridded climate data with species location data to identify climate variables that best predicted the current bioclimatic niches of beaked hazelnut (Corylus cornuta), Oregon grape (Mahonia aquifolium), and salal (Gaultheria shallon). We also developed thermal-sum models for the timing of flowering and fruit ripening for these species. We then used multi-model ensemble future climate projections to estimate how species range and phenology may change under future conditions. Modelling efforts showed extreme minimum temperature, climate moisture deficit, and mean summer precipitation were predictive of climatic suitability across all three species. Future bioclimatic niche models project substantial reductions in habitat suitability across the lower elevation and southern portions of the species' current ranges by the end of the 21st century. Thermal-sum phenology models for these species indicate that flowering and the ripening of fruits and nuts will advance an average of 25 days by the mid-21st century, and 36 days by the late-21st century under a high emissions scenario (RCP 8.5). Future changes in the climatic niche and phenology of these important food-producing species may alter trophic relationships, with cascading impacts on regional ecosystems.

Effectiveness of winter temperatures for satisfying chilling requirements for reproductive budburst of red alder (Alnus rubra).

BACKGROUND: Experiencing an adequate amount of cold temperatures over winter is necessary for many temperate tree species to break dormancy and flower in spring. Thus, changes in winter and spring temperatures associated with climate change may influence when trees break dormancy and flower in the future. There have been several experimental studies that have quantified the effectiveness of cold temperatures for chilling requirements for vegetative budburst of temperate trees; however, there are few experimental studies addressing the chilling requirements for reproductive budburst of trees, as it is difficult to place reproductively mature trees in temperature-controlled environments. METHODS: To identify how changing temperatures associated with climate change may impact reproductive phenology, we completed a temperature-controlled growth chamber experiment using cuttings of reproductive branches of red alder (Alnus rubra), one of the most widespread hardwood tree species of the Pacific Northwest, USA. The purpose of this study was to examine how colder (4 °C) and warmer (9 °C) winter temperature regimes influenced the timing of reproductive budburst of red alder cuttings in spring. We also compared the date of budburst of cuttings to that of branches from intact trees. RESULTS: We found that cuttings flowered earlier after pretreatment with a 4 °C winter temperature regime than after a 9 °C winter temperature regime. We found no significant differences between the timing of male budburst of cuttings exposed to ambient conditions compared to male budburst of branches from intact trees. We used our experimental data to estimate a "possibility-line" that shows the accumulated chilling and forcing temperatures necessary prior to reproductive budburst of red alder. DISCUSSION: This study provides a preliminary indication that warmer winters with climate change may not be as effective as colder winters for satisfying chilling temperature requirements of a Northwest hardwood tree species.

Photoperiod cues and patterns of genetic variation limit phenological responses to climate change in warm parts of species' range: Modeling diameter-growth cessation in coast Douglas-fir.

The phenology of diameter-growth cessation in trees will likely play a key role in mediating species and ecosystem responses to climate change. A common expectation is that warming will delay cessation, but the environmental and genetic influences on this process are poorly understood. We modeled the effects of temperature, photoperiod, and seed-source climate on diameter-growth-cessation timing in coast Douglas-fir (an ecologically and economically vital tree) using high-frequency growth measurements across broad environmental gradients for a range of genotypes from different seed sources. Our model suggests that cool temperatures or short photoperiods can induce cessation in autumn. At cool locations (high latitude and elevation), cessation seems to be induced primarily by low temperatures in early autumn (under relatively long photoperiods), so warming will likely delay cessation and extend the growing season. But at warm locations (low latitude or elevation), cessation seems to be induced primarily by short photoperiods later in autumn, so warming will likely lead to only slight extensions of the growing season, reflecting photoperiod limitations on phenological shifts. Trees from seed sources experiencing frequent frosts in autumn or early winter tended to cease growth earlier in the autumn, potentially as an adaptation to avoid frost. Thus, gene flow into populations in warm locations with little frost will likely have limited potential to delay mean cessation dates because these populations already cease growth relatively late. In addition, data from an abnormal heat wave suggested that very high temperatures during long photoperiods in early summer might also induce cessation. Climate change could make these conditions more common in warm locations, leading to much earlier cessation. Thus, photoperiod cues, patterns of genetic variation, and summer heat waves could limit the capacity of coast Douglas-fir to extend its growing season in response to climate change in the warm parts of its range.

Emerging climate-driven disturbance processes: widespread mortality associated with snow-to-rain transitions across 10° of latitude and half the range of a climate-threatened conifer.

Climate change is causing rapid changes to forest disturbance regimes worldwide. While the consequences of climate change for existing disturbance processes, like fires, are relatively well studied, emerging drivers of disturbance such as snow loss and subsequent mortality are much less documented. As the climate warms, a transition from winter snow to rain in high latitudes will cause significant changes in environmental conditions such as soil temperatures, historically buffered by snow cover. The Pacific coast of North America is an excellent test case, as mean winter temperatures are currently at the snow-rain threshold and have been warming for approximately 100 years post-Little Ice Age. Increased mortality in a widespread tree species in the region has been linked to warmer winters and snow loss. Here, we present the first high-resolution range map of this climate-sensitive species, Callitropsis nootkatensis (yellow-cedar), and document the magnitude and location of observed mortality across Canada and the United States. Snow cover loss related mortality spans approximately 10° latitude (half the native range of the species) and 7% of the overall species range and appears linked to this snow-rain transition across its range. Mortality is commonly >70% of basal area in affected areas, and more common where mean winter temperatures is at or above the snow-rain threshold (>0 °C mean winter temperature). Approximately 50% of areas with a currently suitable climate for the species (<-2 °C) are expected to warm beyond that threshold by the late 21st century. Regardless of climate change scenario, little of the range which is expected to remain suitable in the future (e.g., a climatic refugia) is in currently protected landscapes (<1-9%). These results are the first documentation of this type of emerging climate disturbance and highlight the difficulties of anticipating novel disturbance processes when planning for conservation and management.

Tolerance to multiple climate stressors: a case study of Douglas-fir drought and cold hardiness.

Drought and freeze events are two of the most common forms of climate extremes which result in tree damage or death, and the frequency and intensity of both stressors may increase with climate change. Few studies have examined natural covariation in stress tolerance traits to cope with multiple stressors among wild plant populations.We assessed the capacity of coastal Douglas-fir (Pseudotsuga menziesii var. menziesii), an ecologically and economically important species in the northwestern USA, to tolerate both drought and cold stress on 35 populations grown in common gardens. We used principal components analysis to combine drought and cold hardiness trait data into generalized stress hardiness traits to model geographic variation in hardiness as a function of climate across the Douglas-fir range.Drought and cold hardiness converged among populations along winter temperature gradients and diverged along summer precipitation gradients. Populations originating in regions with cold winters had relatively high tolerance to both drought and cold stress, which is likely due to overlapping adaptations for coping with winter desiccation. Populations from regions with dry summers had increased drought hardiness but reduced cold hardiness, suggesting a trade-off in tolerance mechanisms.Our findings highlight the necessity to look beyond bivariate trait-climate relationships and instead consider multiple traits and climate variables to effectively model and manage for the impacts of climate change on widespread species.

Will changes in phenology track climate change? A study of growth initiation timing in coast Douglas-fir.

Under climate change, the reduction of frost risk, onset of warm temperatures and depletion of soil moisture are all likely to occur earlier in the year in many temperate regions. The resilience of tree species will depend on their ability to track these changes in climate with shifts in phenology that lead to earlier growth initiation in the spring. Exposure to warm temperatures ('forcing') typically triggers growth initiation, but many trees also require exposure to cool temperatures ('chilling') while dormant to readily initiate growth in the spring. If warming increases forcing and decreases chilling, climate change could maintain, advance or delay growth initiation phenology relative to the onset of favorable conditions. We modeled the timing of height- and diameter-growth initiation in coast Douglas-fir (an ecologically and economically vital tree in western North America) to determine whether changes in phenology are likely to track changes in climate using data from field-based and controlled-environment studies, which included conditions warmer than those currently experienced in the tree's range. For high latitude and elevation portions of the tree's range, our models predicted that warming will lead to earlier growth initiation and allow trees to track changes in the onset of the warm but still moist conditions that favor growth, generally without substantially greater exposure to frost. In contrast, toward lower latitude and elevation range limits, the models predicted that warming will lead to delayed growth initiation relative to changes in climate due to reduced chilling, with trees failing to capture favorable conditions in the earlier parts of the spring. This maladaptive response to climate change was more prevalent for diameter-growth initiation than height-growth initiation. The decoupling of growth initiation with the onset of favorable climatic conditions could reduce the resilience of coast Douglas-fir to climate change at the warm edges of its distribution.

A new model to simulate climate-change impacts on forest succession for local land management.

We developed a new climate-sensitive vegetation state-and-transition simulation model (CV-STSM) to simulate future vegetation at a fine spatial grain commensurate with the scales of human land-use decisions, and under the joint influences of changing climate, site productivity, and disturbance. CV-STSM integrates outputs from four different modeling systems. Successional changes in tree species composition and stand structure were represented as transition probabilities and organized into a state-and-transition simulation model. States were characterized based on assessments of both current vegetation and of projected future vegetation from a dynamic global vegetation model (DGVM). State definitions included sufficient detail to support the integration of CV-STSM with an agent-based model of land-use decisions and a mechanistic model of fire behavior and spread. Transition probabilities were parameterized using output from a stand biometric model run across a wide range of site productivities. Biogeographic and biogeochemical projections from the DGVM were used to adjust the transition probabilities to account for the impacts of climate change on site productivity and potential vegetation type. We conducted experimental simulations in the Willamette Valley, Oregon, USA. Our simulation landscape incorporated detailed new assessments of critically imperiled Oregon white oak (Quercus garryana) savanna and prairie habitats among the suite of existing and future vegetation types. The experimental design fully crossed four future climate scenarios with three disturbance scenarios. CV-STSM showed strong interactions between climate and disturbance scenarios. All disturbance scenarios increased the abundance of oak savanna habitat, but an interaction between the most intense disturbance and climate-change scenarios also increased the abundance of subtropical tree species. Even so, subtropical tree species were far less abundant at the end of simulations in CV-STSM than in the dynamic global vegetation model simulations. Our results indicate that dynamic global vegetation models may overestimate future rates of vegetation change, especially in the absence of stand-replacing disturbances. Modeling tools such as CV-STSM that simulate rates and direction of vegetation change affected by interactions and feedbacks between climate and land-use change can help policy makers, land managers, and society as a whole develop effective plans to adapt to rapidly changing climate.

Impact of climate change on cold hardiness of Douglas-fir (Pseudotsuga menziesii): environmental and genetic considerations.

The success of conifers over much of the world's terrestrial surface is largely attributable to their tolerance to cold stress (i.e., cold hardiness). Due to an increase in climate variability, climate change may reduce conifer cold hardiness, which in turn could impact ecosystem functioning and productivity in conifer-dominated forests. The expression of cold hardiness is a product of environmental cues (E), genetic differentiation (G), and their interaction (G × E), although few studies have considered all components together. To better understand and manage for the impacts of climate change on conifer cold hardiness, we conducted a common garden experiment replicated in three test environments (cool, moderate, and warm) using 35 populations of coast Douglas-fir (Pseudotsuga menziesii var. menziesii) to test the hypotheses: (i) cool-temperature cues in fall are necessary to trigger cold hardening, (ii) there is large genetic variation among populations in cold hardiness that can be predicted from seed-source climate variables, (iii) observed differences among populations in cold hardiness in situ are dependent on effective environmental cues, and (iv) movement of seed sources from warmer to cooler climates will increase risk to cold injury. During fall 2012, we visually assessed cold damage of bud, needle, and stem tissues following artificial freeze tests. Cool-temperature cues (e.g., degree hours below 2 °C) at the test sites were associated with cold hardening, which were minimal at the moderate test site owing to mild fall temperatures. Populations differed 3-fold in cold hardiness, with winter minimum temperatures and fall frost dates as strong seed-source climate predictors of cold hardiness, and with summer temperatures and aridity as secondary predictors. Seed-source movement resulted in only modest increases in cold damage. Our findings indicate that increased fall temperatures delay cold hardening, warmer/drier summers confer a degree of cold hardiness, and seed-source movement from warmer to cooler climates may be a viable option for adapting coniferous forest to future climate.

Tradeoffs between chilling and forcing in satisfying dormancy requirements for Pacific Northwest tree species.

Many temperate and boreal tree species have a chilling requirement, that is, they need to experience cold temperatures during fall and winter to burst bud normally in the spring. Results from trials with 11 Pacific Northwest tree species are consistent with the concept that plants can accumulate both chilling and forcing units simultaneously during the dormant season and they exhibit a tradeoff between amount of forcing and chilling. That is, the parallel model of chilling and forcing was effective in predicting budburst and well chilled plants require less forcing for bud burst than plants which have received less chilling. Genotypes differed in the shape of the possibility line which describes the quantitative tradeoff between chilling and forcing units. Plants which have an obligate chilling requirement (Douglas-fir, western hemlock, western larch, pines, and true firs) and received no or very low levels of chilling did not burst bud normally even with long photoperiods. Pacific madrone and western redcedar benefited from chilling in terms of requiring less forcing to promote bud burst but many plants burst bud normally without chilling. Equations predicting budburst were developed for each species in our trials for a portion of western North America under current climatic conditions and for 2080. Mean winter temperature was predicted to increase 3.2-5.5°C and this change resulted in earlier predicted budburst for Douglas-fir throughout much of our study area (up to 74 days earlier) but later budburst in some southern portions of its current range (up to 48 days later) as insufficient chilling is predicted to occur. Other species all had earlier predicted dates of budburst by 2080 than currently. Recent warming trends have resulted in earlier budburst for some woody plant species; however, the substantial winter warming predicted by some climate models will reduce future chilling in some locations such that budburst will not consistently occur earlier.

Climate-related genetic variation in drought-resistance of Douglas-fir (Pseudotsuga menziesii).

There is a general assumption that intraspecific populations originating from relatively arid climates will be better adapted to cope with the expected increase in drought from climate change. For ecologically and economically important species, more comprehensive, genecological studies that utilize large distributions of populations and direct measures of traits associated with drought-resistance are needed to empirically support this assumption because of the implications for the natural or assisted regeneration of species. We conducted a space-for-time substitution, common garden experiment with 35 populations of coast Douglas-fir (Pseudotsuga menziesii var. menziesii) growing at three test sites with distinct summer temperature and precipitation (referred to as 'cool/moist', 'moderate', or 'warm/dry') to test the hypotheses that (i) there is large genetic variation among populations and regions in traits associated with drought-resistance, (ii) the patterns of genetic variation are related to the native source-climate of each population, in particular with summer temperature and precipitation, (iii) the differences among populations and relationships with climate are stronger at the warm/dry test site owing to greater expression of drought-resistance traits (i.e., a genotype × environment interaction). During midsummer 2012, we measured the rate of water loss after stomatal closure (transpiration(min)), water deficit (% below turgid saturation), and specific leaf area (SLA, cm(2) g(-1)) on new growth of sapling branches. There was significant genetic variation in all plant traits, with populations originating from warmer and drier climates having greater drought-resistance (i.e., lower transpiration(min), water deficit and SLA), but these trends were most clearly expressed only at the warm/dry test site. Contrary to expectations, populations from cooler climates also had greater drought-resistance across all test sites. Multiple regression analysis indicated that Douglas-fir populations from regions with relatively cool winters and arid summers may be most adapted to cope with drought conditions that are expected in the future.

Growth phenology of coast Douglas-fir seed sources planted in diverse environments.

The timing of periodic life cycle events in plants (phenology) is an important factor determining how species and populations will react to climate change. We evaluated annual patterns of basal-area and height growth of coast Douglas-fir (Pseudotusga menziesii var. menziesii (Mirb.) Franco) seedlings from four seed sources that were planted in four diverse environments as part of the Douglas-fir Seed-Source Movement Trial. Stem diameters and heights were measured periodically during the 2010 growing season on 16 open-pollinated families at each study installation. Stem diameters were measured on a subset of trees with electronic dendrometers during the 2010 and 2011 growing seasons. Trees from the four seed sources differed in phenology metrics that described the timing of basal-area and height-growth initiation, growth cessation and growth rates. Differences in the height-growth metrics were generally larger than differences in the basal-area growth metrics and differences among installations were larger than differences among seed sources, highlighting the importance of environmental signals on growth phenology. Variations in the height- and basal-area growth metrics were correlated with different aspects of the seed-source environments: precipitation in the case of height growth and minimum temperature in the case of basal-area growth. The detailed dendrometer measurements revealed differences in growth patterns between seed sources during distinct periods in the growing season. Our results indicate that multiple aspects of growth phenology should be considered along with other traits when evaluating adaptation of populations to future climates.

Temperature regulation of bud-burst phenology within and among years in a young Douglas-fir (Pseudotsuga menziesii) plantation in western Washington, USA.

Past research has established that terminal buds of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings from many seed sources have a chilling requirement of about 1200 h at 0-5 degrees C; once chilled, temperatures > 5 degrees C force bud burst via accumulation of heat units. We tested this sequential bud-burst model in the field to determine whether terminal buds of trees in cooler microsites, which receive less heat forcing, develop more slowly than those in warmer microsites. For three years we monitored terminal bud development in young saplings as well as soil and air temperatures on large, replicated plots in a harvest unit; plots differed in microclimate based on amount of harvest residue and shade from neighboring stands. In two of three years, trees on cooler microsites broke bud 2 to 4 days earlier than those on warmer microsites, despite receiving less heat forcing from March to May each year. A simple sequential model did not predict cooler sites having earlier bud burst nor did it correctly predict the order of bud burst across the three years. We modified the basic heat-forcing model to initialize, or reset to zero, the accumulation of heat units whenever significant freezing temperature events (> or = 3 degree-hours day(-1) < 0 degrees C) occurred; this modified model correctly predicted the sequence of bud burst across years. Soil temperature alone or in combination with air temperature did not improve our predictions of bud burst. Past models of bud burst have relied heavily on data from controlled experiments with simple temperature patterns; analysis of more variable temperature patterns from our 3-year field trial, however, indicated that simple models of bud burst are inaccurate. More complex models that incorporate chilling hours, heat forcing, photoperiod and the occurrence of freeze events in the spring may be needed to predict effects of future silvicultural treatments as well to interpret the implications of climate-change scenarios. Developing and testing new models will require data from both field and controlled-environment experiments.

Effects of irrigation, spacing and fertilization on flowering and growth in young Alnus rubra.

Flowering and vegetative growth were assessed in 19 half-sib families of Alnus rubra Bong. planted in a replicated field trial near Olympia, Washington, USA. The trial consisted of three square spacings (0.5, 1.0 and 2.0 m), two irrigation regimes (low and high), and two fertilization treatments (0 and 300 kg P ha(-1)). Male and female flowers were surveyed in all plots for all families at plantation ages 4 and 5 years. Female strobili were surveyed for seven families in the 2-m spaced plots at plantation age 6 years. The percentage of trees flowering and the number of flowers per tree were always greatest, and height and diameter growth were always least, in the low-irrigation regime. Phosphorus fertilization had no effect on the percentage of trees flowering or on 5-year height or diameter growth; it had a positive but small effect on the number of female flowers per tree at age 5 years. Wider spacing resulted in larger trees, higher rates of flowering, and higher tree survival. Within each irrigation regime, the percentage of trees flowering increased as tree size increased. There was substantial variation in flowering among families, with positive but low correlations between tree size and flowering attributes. At ages 4 and 5 years, the ratio of number of trees flowering in the low-irrigation regime to number of trees flowering in the high-irrigation regime differed among families. By age 6 years, many more trees flowered than in previous years, and differences between irrigation regimes were reduced. Early growth rates were rapid and resulted in substantial crown recession and mortality in the closer spacings by age 5 years. We conclude that spacings less than 2 m should only be used in seed production areas if roguing can be done by age 2 to 3 years.