Abstract1. Predictions of upslope range shifts for tree species with warming are based on assumptions of moisture stress at lower elevation limits and low-temperature stress at high-elevation limits. However, recent studies have shown that warming can reduce tree seedling establishment across the entire gradient from subalpine forest to alpine via moisture limitation. Warming effects also vary with species, potentially resulting in community shifts in high-elevation forests.2. We examined the growth and physiology underlying effects of warming on seedling demographic patterns. We evaluated dry mass (DM), root length, allocation above-and below-ground, and relative growth rate (RGR) of whole seedlings, and their ability to avoid or endure water stress via water-use efficiency and resisting turgor loss, for Pinus flexilis, Picea engelmannii and Pinus contorta seeded below, at and above tree line in experimentally warmed, watered and control plots in the Rocky Mountains, USA. We expected that growth and allocation responses to warming would relate to moisture status and that variation in drought tolerance traits would explain species differences in survival rates.3. Across treatments and elevations, seedlings of all species had weak turgor-loss resistance, and growth was marginal with negative RGR in the first growth phase (−0.01 to −0.04 g g −1 day −1 ). Growth was correlated with soil moisture, particularly in the relatively small-seeded P. contorta and P. engelmannii. Pinus flexilis, known to have the highest survivorship, attained the greatest DM and longest root but was also the slowest growing and most water-use efficient. This was likely due to its greater reliance on seed reserves. Seedlings developed 15% less total DM, 25% less root DM and 11% shorter roots in heated compared with unheated plots. Higher temperatures slightly increased DM, root length and RGR where soils were wettest, but more strongly decreased these variables under drier conditions.

Application of stable isotopes of water to studies of plant-soil interactions often requires a substantial preparatory step of extracting water from samples without fractionating isotopes. Online heating is an emerging approach for this need, but is relatively untested and major questions of how to best deliver standards and assess interference by organics have not been evaluated. We examined these issues in our application of measuring woody stem xylem of sagebrush using a Picarro laser spectrometer with online induction heating. We determined (1) effects of cryogenic compared to induction-heating extraction, (2) effects of delivery of standards on filter media compared to on woody stem sections, and (3) spectral interference from organic compounds for these approaches (and developed a technique to do so). Our results suggest that matching sample and standard media improves accuracy, but that isotopic values differ with the extraction method in ways that are not due to spectral interference from organics.

Red spruce (Picea rubens Sarg.) winter injury is caused by freezing damage that results in the abscission of the most recent foliar age-class. Injury was widespread and severe in the northeastern United States in 2003 and was assessed at multiple elevations at 23 sites in Vermont and adjacent states. This paper presents a spatial analysis of these injury assessments. Relationships between winter injury on dominant and codominant spruce trees and elevation, latitude, longitude, slope, and aspect were investigated with least squares regression and geographically weighted regression. Results of these analyses indicate that injury increased (1) with elevation; (2) from east to west; (3) with the degree to which plots faced west, except at the highest elevations, where injury was uniformly severe; (4) with increases in slope steepness at higher elevations, or with decreases in slope steepness at lower elevations; and (5) with the degree to which plots faced south, except at the highest elevations in northern locations, where injury was uniformly severe. Because injury was greater in areas that have historically received higher levels of acid and nitrogen deposition -western portions of the study region, west-facing slopes, and higher elevations -observed patterns of injury support the hypothesis that acidic and (or) nitrogen deposition act on a landscape scale to exacerbate winter injury. Greater injury on south-facing slopes suggests that sun exposure exacerbates injury or its expression. Résumé :Les dommages causés par le froid chez l'épinette rouge (Picea rubens Sarg.) sont dus au gel qui entraîne l'abscission de la classe de feuillage la plus récente. Les dommages étaient largement répandus et sévères dans le nordest des États-Unis en 2003 et ont été évalués à plusieurs altitudes dans 23 sites situés au Vermont et dans les États adjacents. Cet article présente une analyse spatiale de l'évaluation de ces dommages. Les relations entre les dommages causés par le froid sur les tiges d'épinette dominantes et codominantes et l'altitude, la latitude, la longitude, la pente et l'exposition ont été étudiées avec la méthode des moindres carrés et la régression géographiquement pondérée. Les ré-sultats de ces analyses indiquent que les dommages augmentent : (1) avec l'altitude; (2) d'est en ouest; (3) plus les parcelles sont exposées à l'ouest, excepté aux plus hautes altitudes où la sévérité des dommages est uniforme; (4) avec l'augmentation de la pente à haute altitude ou avec la diminution de la pente à basse altitude; et (5) plus les parcelles sont exposées au sud, excepté aux plus hautes altitudes dans les endroits situés au nord où la sévérité des dommages était plus uniforme. Parce que les dommages étaient plus importants dans les zones qui ont historiquement reçu des quantités plus élevées de dépôts acides et d'azote, soit la partie ouest de la région étudiée, les pentes exposées à l'ouest et les secteurs situés en altitude, les patrons de dommages que les auteurs ont observés supportent l'hypothèse que les dépôts ac...

Abundant winter injury to the current-year (2002) foliage of red spruce (Picea rubens Sarg.) became apparent in the northeastern United States in late winter of 2003. To assess the severity and extent of this damage, we measured foliar winter injury at 28 locations in Vermont and surrounding states and bud mortality at a subset of these sites. Ninety percent of all trees assessed showed some winter injury, and trees lost an average of 46% of all current-year foliage. An average of 32% of buds formed in 2002 were killed in association with winter injury. Both foliar and bud mortality increased with elevation and with crown dominance, and bud mortality increased with greater foliar injury. Foliar injury in 2003 at a plantation near Colebrook, New Hampshire, was more than five times the typical levels for 9 previous years of measurement and more than twice that measured for another high-injury year. Plantation data also indicated that bud mortality in 2003 was greater than previously documented and that persistent winter injury was associated with increased tree mortality. Comparisons of our data with past studies for two sites with native red spruce also indicated that damage in 2003 was greater than other recently reported, high-injury years. Because heavy foliar and bud losses can severely disrupt the carbon economies of trees, the 2003 winter injury event could lead to further spruce decline and mortality, particularly among dominant trees at higher elevations.

Where serpentine soils exist, variation in soil properties affects plant species distribution at both coarse and fine spatial scales. The New Idria (California, USA) serpentine mass has barren areas, supporting only sparse shrub and tree islands, adjacent to areas of densely-vegetated serpentine chaparral. To identify factors limiting growth on barren relative to vegetated serpentine soils, we analyzed soils from barren, shrub-island within barren, and vegetated areas and foliage from shrubisland and vegetated areas. We also grew Ceanothus cuneatus (native evergreen shrub), Achillea millefolium (native perennial forb), and Bromus madritensis ssp. rubens (invasive annual grass) in soils from barren and vegetated areas amended factorially with N, K, and Ca in a pot study. In well-watered pots, biomass was greater by 5-, 14-, and 33-fold for Ceanothus, Achillea, and Bromus, respectively, on vegetated-area-collected soils than on barren-collected soils, indicating a strong soil chemistry effect. Although field soil data suggested nutrient deficiency and not heavy metal toxicity, pot study plant data indicated otherwise for two of the three species. On barren-collected soils, only Ceanothus responded positively to added N and Ca and did not show greater foliar Mg or heavy metal (Fe, Ni, Cr, Co, Zn) concentrations than on vegetated-area-collected soils. Ceanothus maintained lower root Mg and heavy metal (Fe, Ni, Cr, Co) concentrations on barren soils and translocated less heavy metal (Fe, Ni, Cr, Co, Mn, Cu) from roots to foliage than Achillea and Bromus. Achillea and Bromus showed significant log-log biomass relationships with foliar Ca:Mg (+), Mg (-), and heavy metals (Fe, Ni, Cr, Co, Mn, Cu, Zn) (-), while Ceanothus showed relationships only with Ca: Mg (+) and Mg (-). The New Idria barren-vegetated pattern appears to be maintained by different factors for different species or functional types-low Ca:Mg ratios on barrens for all species tested, high heavy metal concentrations for Achillea and Bromus, and low macronutrient (N) concentrations for Ceanothus. Combined data from this and other studies suggest high heavy metal concentrations more strongly affect Plant Soil (2011) herbaceous than woody species, contributing to variation in species distribution on serpentine soils.

1. Despite broad recognition that water is a major limiting factor in arid ecosystems, we lack an empirical understanding of how this resource is shared and distributed among neighbouring plants. Intraspecific variability can further contribute to this variation via divergent life-history traits, including root architecture. We investigated these questions in the shrub Artemisia tridentata and hypothesized that the ability to access and utilize surface water varies among subspecies and cytotypes. We used an isotope tracer to quantify below-ground zone of influence inA. tridentata, and tested whether spatial neighbourhood characteristics can alter plant water uptake. We introduced deuterium-enriched water to the soil in plant interspaces in a common garden experiment and measured deuterium composition of plant stems. We then applied spatially explicit models to test for differential water uptake by A. tridentata, including intermingled populations of three subspecies and two ploidy levels.3. The results suggest that lateral root functioning in A. tridentata is associated with intraspecific identity and ploidy level. Subspecies adapted to habitats with deep soils generally had a smaller horizontal reach, and polyploid cytotypes were associated with greater water uptake compared to their diploid variants. We also found that plant crown volume was a weak predictor of water uptake, and that neighbourhood crowding had no discernable effect on water uptake. 4. Intraspecific variation in lateral root functioning can lead to differential patterns of resource acquisition, an essential process in arid ecosystems in the contexts of changing climate and seasonal patterns of precipitation. Altogether, we found that lateral root development and activity are more strongly related to genetic variability within A. tridentata than to plant size. Our study highlights how intraspecific variation in life strategies is linked to mechanisms of resource acquisition. K E Y W O R D SArtemisia tridentata (big sagebrush), below-ground interactions, deuterium tracer, intraspecific variation, lateral roots, resource competition, surface water uptake | 1171 Functional Ecology ZAIATS eT Al.

Premise Physiological responses to temperature extremes are considered strong drivers of species’ demographic responses to climate variability. Plants are typically classified as either avoiders or tolerators in their freezing‐resistance mechanism, but a gradient of physiological‐threshold freezing responses may exist among individuals of a species. Moreover, adaptive significance of physiological freezing responses is poorly characterized, particularly under warming conditions that relax selection on cold hardiness. Methods Freezing responses were measured in winter and again for new foliage in spring for 14 populations of Artemisia tridentata collected throughout its range and planted in a warm common garden. The relationships of the freezing responses to survival were evaluated in the warm garden and in two colder gardens. Results Winter and spring freezing resistance were not correlated and appeared to be under differing selection regimes, as evident in correlations with different population climate of origin variables. All populations resisted considerably lower temperatures in winter than in spring, with populations from more continental climates showing narrower freezing safety margins (difference in temperatures at which ice‐nucleation occurs and 50% reduction in chlorophyll fluorescence occurs) in spring. Populations with greater winter freezing resistance had lower survivorship in the warmest garden, while populations with greater spring freezing resistance had lower survivorship in a colder garden. Conclusions These survivorship patterns relative to physiological thresholds suggest excess freezing resistance may incur a survival cost that likely relates to a trade‐off between carbon gain and freezing resistance during critical periods of moisture availability. This cost has implications for seed moved from cooler to warmer environments and for plants growing in warming environments.

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